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Full text of "Bulletin of the Natural History Museum Entomology"

ISSN 0968-0454 



Bulletin of 
The Natural History Museum 



Entomology Series 




VOLUME 63 NUMBER 2 24 NOVEMBER 1994 



The Bulletin of The Natural History Museum (formerly: Bulletin of the British 
Museum (Natural History) ), instituted in 1949, is issued in four scientific series, 
Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. 

The Entomology Series is produced under the editorship of the 

Keeper of Entomology: Dr R.P. Lane 

Editor of Bulletin: Dr P.C. Barnard 



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World List abbreviation: Bull. nat. Hist. Mus. Lond. (Ent.) 

© The Natural History Museum, 1994 

Entomology Series 
ISSN 0968-0454 Vol. 63, No. 2, pp. 137-210 

The Natural History Museum 

Cromwell Road 

London SW7 5BD Issued 24 November 1994 

Typeset by Ann Buchan (Typesetters), Middlesex 
Printed in Great Britain at The Alden Press, Oxford 



Bull. Br. nat. Hist. (Ent.) 63(2): 137-210 



Issued 24 November 1994 



A taxonomic review of the common 
green lacewing genus Chrysoperla 
(Neuroptera: Chrysopidae) 

S.J. BROOKS 

Department of Entomology, The Natural History Museum, Cromwell Road, 
London SW7 5 BD 

CONTENTS 



Introduction 137 

Historical review 138 

Generic classification 1 38 

Species classification 1 39 

Materials and methods 140 

Abbreviations 1 40 

Acknowledgements 141 

The systematic position of Chrysoperla 141 

Species affinities within Chrysoperla 142 

Chrysoperla Steinmann 143 

Key to the world Chrysoperla species 144 

Species descriptions 146 

Synoptic checklist of the species of Chrysoperla 173 

References 1 74 

Index 210 



Synopsis. The genus Chrysoperla Steinmann (Neuroptera: Chrysopidae), world-wide 
distribution, is revised. Several species of the genus, and in particular Chrysoperla 
carnea (Stephens), are cultured extensively for use in biological control programmes but 
until now the systematics of the genus has remained confused. 36 species are recognized 
as valid. Five new species are described, 4 species have revised status and there are 40 
new specific synonyms, one new provisional synonym and four lectotype designations. 
Eight species are newly excluded from Chrysoperla. A key to the species is provided. 
Male and female of each species are described and diagnostic features figured. 



INTRODUCTION 



More effort has been spent on research into the 
ecology and biology of Chrysoperla than all the 
rest of the chrysopid genera put together (see 
Canard et al., 1984). This is chiefly because 
several species, but especially C. carnea 
(Stephens), have proved easy to culture and their 
larvae are fierce predators of aphids and coccids. 
C. carnea and some of its congeners are now 
commonly reared in biological control laborato- 



revision of the 
Barnard, 1990), 
defined and it is 



ries all over the world. One aspect that has been 
largely overlooked, however, is the systematics 
of the genus as a whole. 

Following a genus-level 
Chrysopidae (Brooks & 
Chrysoperla is now clearly 
possible to correctly assign species to the genus. 
As currently recognized, the genus includes 36 
species, distributed throughout the world but 
with a preponderance of Holarctic species. 
Although it is relatively easy to recognize which 
species belong to the genus, species level identifi- 



)The Natural History Museum. 1994 



138 



S.J. BROOKS 



cation is less straight forward. Tjeder (1966) 
provided a key to five species from southern 
Africa and Holzel (1989b) gave descriptions of a 
further five Afrotropical species. Three western 
European species can be identified using the keys 
in Aspock et al (1980). In order to identify the 
remaining Old World species it is usually neces- 
sary to compare the frequently inadequate origi- 
nal descriptions that are scattered throughout the 
literature often in old and obscure volumes. The 
New World also has a rich Chrysoperla fauna but 
similar identification problems will be encoun- 
tered as there are no modern keys available to 
identify the adults, although Tauber (1974) pro- 
vided descriptions of the North American larvae. 

A further complication when trying to estab- 
lish the correct name for a particular Chrysoperla 
specimen is that, although a few identification 
keys are available for particular regions, regional 
faunas have never been compared with each 
other. This has undoubtedly led to synonymy and 
this study shows that some species occur in more 
than one zoogeographic region where they were 
referred to under different names. 

In recent years traditional morphological taxo- 
nomic studies of Chrysoperla species, especially 
those in the C. carnea species-complex, have 
been supplemented by investigations employing 
additional techniques. One example is the pio- 
neering work begun by Henry (1979) who stud- 
ied the production of substrate-borne calls by 
males and females during courtship. This work 
has necessitated a reappraisal of species limits 
because morphologically identical populations 
have been shown to be reproductively isolated by 
their different courtship songs (Henry, 1983). A 
useful summary of this work appears in Henry 
(1992). Other new approaches include multivari- 
ate analysis of morphological characters (Thierry 
et al., 1992), morphometric studies of the male 
genitalia (Semeria, 1992) and electrophoretic 
studies (Cianchi & Bullini, 1992). These and 
other studies on the Chrysoperla carnea-complex 
are discussed by Thierry & Adams (1992). 

These recent investigations have produced 
some fascinating results and point up some of the 
limitations of a traditional morphological taxo- 
nomic study. However, a thorough revision of 
the genus on a worldwide basis is an essential 
prerequisite if we are to interpret the phyloge- 
netic relationships of the species within the 
genus. Such a revision will also establish the 
geographic distribution of the morpho-species 
and the extent of geographical variation within 
taxa. The new techniques described above have 
only been applied to a few populations in western 
Europe and North America where a plentiful 



supply of material is readily available. In order to 
conduct a comprehensive revision of the world 
species it is necessary to fall back on a morpho- 
logical study of long-dead material stored in 
museum collections. 

This study is the first revision of the world 
species of this economically important genus. As 
a result of this work 36 species are now recog- 
nized as valid and these are described and fig- 
ured. Five new species are described, 4 species 
have new status, 40 new specific synonyms, three 
revised synonyms and one provisional synonym 
are proposed. The aim of this work is to provide 
an overview of the species of Chrysoperla and a 
baseline for future taxonomic work, using a 
wider range of techniques. 

In general, I have decided to recognise as 
species those taxa that have clearly defined, 
unambiguous, morphological characters. Such 
characters are usually derived from features of 
the male genitalia but include other characters 
too. Geographical variation within each species 
is described. Such variation may be indicative of 
sibling species that may be more clearly revealed 
using some of the techniques described above. A 
few species have recently been described that are 
distinguishable only by the courtship songs that 
they produce or by certain aspects of their ecol- 
ogy. I have not synonymised these taxa since the 
biological evidence is compelling and active 
research is still in progress. 



HISTORICAL REVIEW 



Generic classification 

Chrysoperla was first proposed by Steinmann 
(1964) as one of four subgenera of Chrysopa 
Leach. The subgenera were based on differences 
in head markings. However, characters such as 
these have proved to be unreliable in defining 
supraspecific monophyletic groups (Brooks & 
Barnard, 1990) and Steinmann's subgenera have 
no validity as he defined them. The systematics 
of Chrysopidae was revolutionized when Tjeder 
(1966) recognized that the presence or absence of 
certain components in the male genitalia united 
groups of species in the otherwise homogeneous 
genus Chrysopa s.l. Until then this large and 
unwieldy taxon included the large majority of 
chrysopid species. Tjeder considered these 
groups of species as subgenera of Chrysopa. One 
of the eight subgenera he recognized included 
those species in which the arcuate dorsal struc- 
ture in the male genitalia, termed the tignum. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



139 



was present but the gonapsis, a ventral winged 
structure, was absent. This species group 
included C. cornea (Stephens) which Steinmann 
had nominated the type-species of Chrysoperla. 
Tjeder (1966) included 17 species in Chrysoperla 
which were divided between two species groups. 
The second of these species groups is now consid- 
ered to be a distinct genus, Atlantochrysa Holzel 
(Brooks & Barnard, 1990). Following Tjeder's 
lead. Holzel (1970) revised the generic classifica- 
tion of the Palaearctic Chrysopinae. Holzel 
accorded full generic status to some of the more 
distinctive taxa previously grouped together in 
Chrysopa s.l. but those groups in which males 
possessed a tignum and/or gonapsis, including 
Chrysoperla, were included together as subgen- 
era of Anisochrysa Nakahara (now Mallada 
Navas). 

Semeria (1977) later raised Chrysoperla to full 
generic status. He showed that species of 
Chrysoperla were distinct from those of Anisoch- 
rysa in three principal ways. First, males of 
Anisochrysa possess a gonapsis, which is absent 
in Chrysoperla, secondly larvae of Anisochrysa 
carry debris but are naked in Chrysoperla and 
finally, the adult is the over-wintering stage in 
Chrysoperla but in Anisochrysa it is the third 
instar larva that over-winters. 

Electrophoretic studies by Bullini et al. (1984) 
supported the recognition of Chrysoperla as dis- 
tinct from other chrysopid genera and moreover 
demonstrated that it was more closely related to 
Anisochrysa (= Mallada) than to Chrysopa 
Leach. 

The first genus-level revision of the world 
Chrysopidae was published by Brooks & Bar- 
nard (1990). In that paper, Chrysoperla was 
demonstrated to be monophyletic, closely related 
to but distinct from Mallada. As well as the 
characters noted by previous workers as distin- 
guishing Chrysoperla, additional autapomorphies 
for the genus were identified. 

Species classification 

Although the species of Chrysoperla are superfi- 
cially rather homogeneous, there is considerable 
variation in the extent and colour of markings, 
especially on the head, within and between taxa. 
Many of the taxa are also common, abundant 
and widely distributed. This combination of fac- 
tors has lead to a plethora of species-group 
names appearing in the literature. During this 
study 112 species-group names assignable to 
Chrysoperla were traced (plus numerous varietal 
names) but I have treated only 36 as valid. 
The wide distributional range and degree of 



morphological variation between populations of 
some Chrysoperla species led some authors to 
doubt whether these taxa are true biological 
entities. In order to tackle this problem some 
workers have adopted additional methods that 
supplement the traditional use of morphological 
characters. These methods include the analysis of 
courtship songs", multivariate analysis of mor- 
phological characters, morphometric methods 
and biochemical techniques. 

Henry (1979) investigated courtship and mat- 
ing in North American examples of C. 
plorabunda and found that rhythmic tremulation 
of the abdomen occurred in both sexes through- 
out courtship. These non-audible vibrations arc 
transmitted through the substrate and picked up 
by receptors in the insects' legs. Further work 
(e.g. Henry. 19X3: 1985a & b; Henry & Wells. 
1990; Henry et al., 1993) has revealed the exist- 
ence of song morphs' that are morphologically 
indistinguishable but produce different courtship 
songs. In some Nearctic localities these song 
morphs occur sympatrically but will not naturally 
interbreed and Henry (1993) has described these 
as distinct sibling species. However, the signifi- 
cance of song differences in at least some 
instances remains unclear. Wells (1991) was able 
to produce fertile hybrids in the laboratory 
between two North American song-morphs (C. 
plorabunda and ('. johnsoni) and Duelli (1987) 
reported that hybridisation did occur in the labo- 
ratory between two song morphs in Switzerland. 
Wells (1991), using gel electrophoresis, has 
found the genetic distance between three song- 
morphs of C. plorabunda to be no greater than 
that usually found between individuals of the 
same animal species. The songs of few European 
populations have been examined but Henry 
(1983) has shown that individuals of C. carnea 
from Switzerland produce songs that are radi- 
cally different from those of otherwise indistin- 
guishable nearctic populations. 

Thierry el al., (1992) have adopted a different 
approach and conducted a multivariate analysis 
of eight morphological character states in French 
populations of C. carnea. Their results indicate 
that there are three distinct adult morphs that 
can be associated with three larval morphs. 
While refraining from naming these morphs as 
distinct species, they suggest that these morphs 
may represent sibling species. 

Detailed morphometric studies of the male 
genitalia by Semeria (1992) have suggested the 
occurrence of six sibling species of Chrysoperla 
carnea in France. 

Electrophoretic and genetic studies of Euro- 
pean populations of C. carnea have also sug- 



140 



S.J. BROOKS 



gested the presence of sympatric sibling species 
(Bullini & Cianchi, 1984; Bullini et at., 1983; 
1984; Cianchi & Bullini, 1992). Results suggest 
that certain central European populations are 
reproductively isolated. However, the values of 
genetic distance (Nei, 1972) are relatively low so 
it is postulated that they have recently diverged. 

The occurrence of sibling species in the C. 
carnea complex has also been identified follow- 
ing ecological studies. Tauber & Tauber (1977 a; 
b) identified two sympatric sibling species within 
Nearctic populations of the cornea-complex . One 
of the sibling species, C. downesi, occurs in 
conifer forests and is dark green all year round. 
The second sibling species, C. plorabunda, 
occurs in meadows during summer, when it is 
bright green, but during the winter is found in 
deciduous woodland and is brown. Morphologi- 
cally these two taxa are otherwise indistinguish- 
able. 

The work described above presents difficulties 
in interpretation especially when the genus as a 
whole is being reviewed. First, only a few taxa 
have been investigated in such ways and then 
from only a limited geographical range. Also 
there has been no integration between these 
techniques. At present none of the so-called 
song-morphs have been examined using multi- 
variate techniques and few have been investi- 
gated using electrophoresis. DNA studies of any 
of the above sibling species may also produce 
useful comparative results. 

Until Chrysoperla populations have been 
investigated more comprehensively I think that it 
is unwise to introduce even more names to an 
already confusing picture. In this work I have 
decided to recognise species if they can be clearly 
defined using classic morphological characters, 
such as those that occur in the male genitalia, or 
if there is overwhelming evidence from other 
techniques. This should provide a sound frame- 
work on which to base further studies on the 
possible occurrence of sibling species. 



examined and in many cases the genitalia dis- 
sected. Drawings were made using a camera 
lucida attachment on a stereo dissecting micro- 
scope or compound microscope. The genitalia 
were stored in glycerol in glass vials or perma- 
nently slide mounted. The type specimens of 
many species were examined but, where this was 
not possible, reliably identified specimens were 
examined instead. 

In the synonymic lists provided with each 
species description only species and subspecies 
are listed. For several species large numbers of 
named varieties have also been described. Most 
of these have not been examined. If this had 
been attempted the length of time spent on the 
project would have been considerably increased 
but almost certainly would have had no impact 
on the validity of the species-group names used 
in this study. 

There are several morphological characters 
which were found to be particularly useful when 
distinguishing species or determining phyloge- 
netic relationships. Some of these involved com- 
parative measurements using a calibrated 
graticule. 

Length of costal setae 

The length of the fringe of setae on the costal 
margin of the fore wing was determined by 
measuring the height of the fringe at right angles 
to the costal vein at the level of the second radial 
crossvein. The more erect the setae, the longer 
the costal fringe will appear. 

Fore wing length: breadth 

This ratio is obtained by dividing the overall 
length of the fore wing by the width at the wing's 
widest point, measuring perpendicularly from 
the costa. 

Basal dilation of tarsal claw 



MATERIALS AND METHODS 



The relative size of the basal dilation is expressed 
as a ratio of the claw hook length to the length of 
the basal dilation of the hind tarsal claw. 



This work has been based primarily on the exten- 
sive collections of Chrysoperla specimens held at 
The Natural History Museum, London, although 
additional specimens have been kindly loaned by 
many institutions and individuals around the 
world. The types of 61 of the 117 described 
species assignable to Chrysoperla have been 
studied and over 2000 specimens have been 



ABBREVIATIONS 



BAU Beijing Agricultural University, Beijing, 

China 
BMNH The Natural History Museum, London, UK 
BPBM Bernice P. Bishop Museum, Honolulu, USA 
CAS California Academy of Sciences, San Fran- 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



141 



cisco, USA 
CNC Canadian National Collection, Ottawa, 

Canada 
HU Hokkaido University, Sapporo, Japan 

IP Institut fur Pflanzenschutzforschung, Eber- 

swalde, Germany 
IRSM Institut de Recherche Scientifique de Mada- 
gascar 
IRSNB Institut Royal des Sciences Naturelles de 

Belgique, Bruxelles, Belgium 
ITZ Instituut voor Taxonomische Zoologie, 

Amsterdam, The Netherlands 
IZAS Institute of Zoology Academia Sinica, 

Beijing, China 
KU Kyushu University, Fukuoka. Japan 

LACM Los Angeles County Museum, Los Angeles, 

USA 
LP La Plata Museum, Buenos Aires, Argentina 

MCSN Museo Civico di Storia Naturale, Genoa, 

Italy 
MCZ Museum of Comparative Zoology, Cam- 
bridge, USA 
MNHN Museum National d'Histoire Naturelle, 

Paris, France 
MRAC Musee Royal de l'Afriquc Centrale, Tcr- 

vuren. Belgium 
NCI National Collection of Insects, Pretoria, 

Republic of South Africa 
NHMZ Natural History Museum of Zimbabwe. 

Bulawayo, Zimbabwe 
NMHU Museum fur Naturkunde der Humboldt- 

Univcrsitat. Berlin, Germany 
NMW Naturhistorisches Museum Wien, Vienna, 

Austria 
NNM Nationaal Natuurhistorisch Museum, 

Leiden. The Netherlands 
NR Naturhistoriska Riksmuseum. Stockholm. 

Sweden 
SDNHM Natural History Museum, San Diego. USA 
TM Transvaal Museum, Pretoria, Republic of 

South Africa 
USP Museum of Zoology, University of Sao 

Paulo, Brazil 
ZMH Zoologisches Museum, Hamburg, Germany 
ZSBS Zoologisches Sammlung des Bayerischen 

Staater, Munich, Germany 
ZSI Zoological Survey of India, Calcutta, India 



Acknowledgements. I would like to thank the follow- 
ing for the loan of specimens: Dr P. A. Adams (Fuller- 
ton University, USA), Dr H.M. Andre (MRAC), Dr 
R. Gaedike (IP), Dr N. Evenhuis (BPBM), Dr D. 
Faulkner (SDNHM), Mrs M.J. Fitzpatrick (NHMZ), 
Dr S. de Freitas (USP), Dr C.T. Graham (MCZ), Dr 
P. Grootaert (IRSNB), Prof. C.S. Henry (University of 
Connecticut, USA), Mr W. Hogenes (ITZ), Herr H. 
Holzel (Bruckl, Austria), Dr Kuldip Rai (ZSI), M J. 
Legrand (MNHN), Dr B. Gustafsson (NR), Dr M.W. 



Mansell (NCI), Dr V.J. Monserrat (University of 
Madrid, Spain), Prof. S. Nomura (KU), Dr N.D. 
Penny (CAS). Mr C.W. Plant (Passmore Edwards 
Museum. UK). Dr R. Poggi (MCSN). Mr R.R. Snel- 
ling (LACM), Dr S. Takagi (HU). Prof. C.A. Taubcr 
(Cornell University, USA), Mr J. van Tol (NNM), Dr 
X.-k. Yang (IZAS). 

I am also grateful to the following for fruitful 
exchange of ideas: Dr P. A. Adams (California State 
University, Fullerton, USA). Prof. C.S. Henry (Uni- 
versity of Connecticutt, Storrs, USA). Prof. E.G. 
MacLeod (University of Illinois, Urbana, USA), Prof. 
C.A. Tauber (Cornell University. Ithaca. New York). 



THE SYSTEMATIC POSITION OF 
CHRYSOPERLA 

Brooks & Barnard (1990) demonstrated that 
Chrysoperla belonged to the Chrysopini which, 
with 30 genera, is the largest tribe of Chrysopi- 
dae. There are a number of adult characters that 
may shed light on the phylogenetic relationships 
of Chrysoperla. 

In discussing the phylogenetic significance of 
male genitalic components in Chrysopidae, 
Brooks & Barnard (1990) concluded that posses- 
sion of a tignum and gonapsis was autapomor- 
phic for the tribe but plesiomorphic within the 
Chrysopini. Because of the morphological uni- 
formity of the tignum throughout the tribe, it is 
unlikely that the tignum has evolved more than 
once but may have been lost on more than one 
occasion. An arcuate tignum occurs in six 
chrysopid genera: Chrysopidia (subgenus Anach- 
rysa) Holzel, Anomalochrysa McLachlan, Atlan- 
tochrysa Holzel, Chrysoperla Steinmann, 
Dichochrysa Yang & Yang and Mallada Navas. 
Like Chrysoperla, Chrysopidia (Anachrysa) and 
Mallada (satilota-group) do not have a gonapsis 
but the gonapsis may have been lost indepen- 
dently in these taxa. A gonapsis is present in the 
other genera mentioned above including the 
Mallada desjardinsi-group. 

The yellow median thoracic band is a charac- 
teristic feature of most Chrysoperla species. 
Within the family this marking is restricted to the 
chrysopine genera Anomalochrysa, Apertochrysa 
Tjeder, Chrysoperla, Chrysopidia (subgenus 
Chrysotropia) Navas, Cunctochrysa Holzel, 
Dichochrysa, Mallada and Peyerimhoffina Lac- 
roix. 

Many species of Chrysoperla possess dorsal 
striations on the arcessus. This character is prob- 
ably derived in Chrysopidae since it does not 
occur elsewhere in the Neuroptera. The occur- 



142 



S.J. BROOKS 



rence of this character in other chrysopid genera 
suggests that they are closely related. There are 
only five other chrysopid genera, and like 
Chrysoperla all are Chrysopini, that have a stri- 
ated arcesuss: Apertochrysa, Atlantochrysa, 
Dichochrysa, Meleoma and Suarius Navas. 

Possibly the most frequently used character for 
distinguishing Chrysoperla in keys is the short 
intramedian cell, the apex of which does not 
usually reach the first crossvein from the radial 
sector. This character, although not occurring 
universally in species of Chrysoperla, may be a 
synapomorphy linking Chrysoperla with two 
other chrysopine genera, Mallada and Peyerim- 
hoffina. The character does not occur elsewhere 
in the Chrysopidae. 

The sister group to Chrysoperla plus the other 
chrysopine genera discussed above is probably 
Dichochrysa. Dichochrysa is linked to these gen- 
era by several synapomorphies but also retains 
some plesiomorphies, particularly in the male 
genitalia, that have become modified in the other 
genera. For this reason, Dichochrysa will be 
considered as the outgroup when deciding on 
character polarities in postulating species rela- 
tionships within Chrysoperla. 



SPECIES AFFINITIES WITHIN 
CHRYSOPERLA 

There are four distinct species groups within the 
genus based on the following adult characters. 

1. carnea-group. This group is the least derived 
and retains many plesiomorphic chrysopine char- 
acters, particularly in the male genitalia. The 
male genitalia of the group lack spinellae, 
gonosetae are generally few, the arcessus is dor- 
sally striated, the entoprocessus are large and 
crescentic, the gonarcus arch is broad and the 
lateral plates are large and reniform, and the 
acumen of the tignum is narrow at its apex. Most 
of these characters are plesiomorphic for the 
Dichochrysa-group of genera, although the large 
crescentic entoprocessus are characteristic of the 
cornea-species group and are probably apomor- 
phic. The carnea-group can be divided into two 
sub-groups. 

1.1. The first sub-group includes the species 
carnea (Palaearctic), furcifera, nipponensis, 
oblita, sillemi (Eastern Palaeartic), iranica 
(Middle East), renoni, mediterranea (Western 
Palaearctic), zastrowi (Afrotropics) and 
plorabunda, adamsi, johnsoni and downesi 
(Nearctic). In these species, with the exception 



of C. oblita, the basal dilation of the claws is 
small (apomorphic for this species group). 

1.2. The second sub-group comprises gallagh- 
eri (Eremic) and annae (Eastern Palaearctic). 
The numerous gonosetae and small, narrow 
entoprocessus in the male genitalia are genus- 
level apomorphies that set this species-subgroup 
apart. 

The following species-groups (2, 3, and 4) all 
possess numerous gonosetae which is apomor- 
phic within Chrysoperla. The possession of a few 
gonosetae, arranged in a lateral group, is a 
synapomorph that links most of the genera in the 
Dichochrysa group. Species groups 2 and 3 are 
linked by the synapomorphic character of broad 
fore wings and long costal setae. 

2. comans-group. This group comprises 
comans, decaryana (both Afrotropics) and sia- 
mensis (Thailand). In these species spinellae are 
absent (plesiomorphic within Chrysoperla), but 
the species share two characters that are apomor- 
phic within the genus: the arcessus is unstriated 
dorsally and the entoprocessus are very small. 
However, this species-group is not well-defined 
since both these apomorphies occur in other 
Chrysoperla species-groups and, in addition, an 
unstriated arcessus is considered to be a second- 
ary character reversal. Nevertheless, the absence 
of spinellae in the male genitalia of group 2 
species preclude their inclusion in groups 3 and 4 
since this is autapomorphic for those groups. 

3. nyerina-group. Species in this group, like 
group 2, also have broad fore wings and long 
costal setae, which is apomorphic within 
Chrysoperla. In addition, males possess spinellae 
on the gonosetae. This character is synapomor- 
phic for groups 3 and 4 and does not occur 
elsewhere in the Chrysopidae. The nyerina- 
group can be divided into two sub-groups. 

3.1. The first sub-group, comprising nyerina 
and plicata (both Afrotropics), is characterised 
by the presence of dorsal striations on the arces- 
sus (plesiomorphic in Chrysoperla) and the pos- 
session of simple spinellae. The spinellae show 
several character states within Chrysoperla. C. 
plicata exhibits the simplest condition in which 
the spinellae arise directly from the gonosaccus. 
In nyerina the spinellae are modified so that each 
spine is positioned on a small tubercle. 

3.2. C. insulata (Reunion Island) is the only 
species in this sub-group. The spinellae are more 
complex than those found in sub-group 3.1, 
being formed from 2-3 spines on a small 
tubercle. This is the only species in the genus 
which lacks a tignum in the male genitalia. This 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



143 



structure has probably been secondarily lost. 

4. pudiea-group. This group is characterised 
by the possession of complex spinellae formed 
from 5 or more spines on each tubercle. This is 
the most derived condition of the spinellae within 
the genus. In this group the fore wings are 
narrow with short costal setae. The shape and 
length of the apex of the arcessus was found to be 
particularly useful in distinguishing species of this 
group. There are two sub-groups. 

4.1. This sub-group includes the species 
pudica (Afrotropics), mutata and nigrinervis 
(Palaearctic). The gonarcus retains its typical 
broad arch configuration with large, reniform 
lateral plates and small entoprocessus. The acu- 
men is swollen apically. At present no apomor- 
phic characters have been found to define this 
sub-group. 

4.2. In the second sub-group most species are 
united by the autapomorphy of a narrow gonar- 
cus arch (which does not occur elsewhere in 
Chrysoperla or in related genera) with narrow 
lateral plates and minute entoprocessus. How- 
ever, dozieri and defreitasi are exceptional in 
retaining a relatively broad gonarcus arch. Spe- 
cies included are: exul (St Helena), congrua (Old 
World tropics), asoralis, exotera, externa, galapa- 
goensis, mexicana (all Nootropics). These species 
all possess an apically swollen acumen. However, 
in the species, comanche, harrisii, rufilabris (all 
Nearctic), defreitsi (Brazil) and dozieri (Haiti), 
the acumen is narrow apically. 

Chrysoperla Steinmann 

Chrysoperla Steinmann, 1964: 260 [as subgenus 
of Chrysopa Leach]. Type-species: Chrysop- 
erla carnea Stephens, 1836, by original desig- 
nation. [As subgenus of Anisochrysa Nakahara 
by Holzel, 1970: 51; raised to genus by Seme- 
ria, 1977: 238.] 

Adult. Fore wing 9-14 mm. Ground colour 
pale green, becoming brown in over-wintering 
specimens. Head sometimes unmarked but usu- 
ally with red or blackish brown stripe on gena 
and lateral clypeus. Maxillary palps unmarked or 
with black dorsal stripe. Labrum indented anteri- 
orly. Mandibles broad, asymmetrical with basal 
tooth on left mandible. Head width : eye width = 
1.8-2.6 : 1. Antenna usually shorter than fore 
wing; flagellar segments 2-3 times as long as 
broad; setae arranged in four rings. Pro-, meso- 
and metanotum usually with yellow or white 
longitudinal median band, sometimes marked 
with lateral red spots; dorsal setae long and pale 
or short and dark. Leg unmarked; claw with, or 



occasionally without, basal dilation. Fore wing 
unmarked; narrow or broad (length : breadth = 
2.6-4.1 : 1); venation green or with gradate 
crossveins black; costal setae short, inclined 
towards wing apex or sometimes long, erect; 
basal costal crossveins sinuous; stigma 
unmarked; cell im narrow, ovate; 1st Rs cross- 
vein usually meeting Psm distad of apex of im, 
but sometimes meets im at apex or subapically; 
gradates in two parallel series, basal inner gra- 
date meets Psm. Abdomen usually with longitu- 
dinal median yellow or white stripe. 

GenitaliacT. Apex of sternite 8+9 with distinct 
lip beset with short setae. Ectoproct with invagi- 
nation ventrad of callus cercus. Gonosaccus with 
gonosetac. with or without spinellae. Arcessus 
narrow with or without dorsal striations. Ento- 
processus crescentic, large or minute. Gonarcus 
broadly arcuate with reniform lateral plates or 
arch narrow with narrow lateral plates. Tignum 
arcuate; acumen long and narrow or short, swol- 
len. Gonapsis absent. 

GENITALIAS'. Praegenitale absent. Subgenitale 
bilobed apically. tapering and extended basally. 
Spermatheca narrow; ventral impression slight: 
vela short; duct short. 

Larva. Abdomen narrow. fusiform. not 
humped. Thoracic and abdominal tubercles 
small, spherical; setae short, smooth, not hooked 
apically: transverse row of metanotal setae 
absent; latero-dorsal chalazae present but indis- 
tinct, bearing single seta. Debris not carried. 

Rt marks Adult males of Chrysoperla are char- 
acterised by the autapomorphic possession of a 
lip at the apex of sternite 8+9. The fusiform 
larvae with small tubercles and non-debris carry- 
ing habit are also distinctive. These characters 
are apomorphic at the genus-level. Other auta- 
pomorphies, but which are not shared by all 
species in the genus, include: the presence of 
spinellae and an over-wintering diapause occur- 
ring in the adult stage. 

As a result of this study it is now apparent that, 
in order to maintain Chrysoperla as a monophyl- 
etic group, a few species listed under Chrysoperla 
by Brooks & Barnard (1990) should be moved to 
different genera. The confusion has arisen 
because the possession of a tignum but lack of a 
gonapsis was previously considered apomorphic 
for Chrysoperla. However, it is now clear that 
the gonapsis has been lost independently at least 
twice in the Chrysopidae. Consequently, the 
species anpingensis Esben-Petersen, australis 
New, krakatauensis Tsukaguchi, maquilingi 



144 



S.J. BROOKS 



Banks, meloui Navas, satilota Banks, suzukii 
Okamoto and triactinata New, in which sternite 
8+9 is not lipped in males, must be reassigned 
(summarised in checklist, p. 173). 

Yang & Yang (1990b) proposed that the Mal- 
lada boninensis (Okamoto) (now a synonym of 
M. desjardinsi Navas (Holzel & Ohm, 1992)) 
-group of species (Brooks & Barnard, 1990) be 
considered as a distinct genus and that the 
remaining Mallada species be included in a new 
genus, Dichochrysa Yang & Yang. This proposal 
has merit since there are several autapomorphies 
that define Mallada s. str., and Mallada s.l. 
seems to be paraphyletic. 

The males of Mallada s. str. are characterised 
by narrow ectoprocts, subapical lobe on sternite 
8+9, two pairs of gonosetae in the genitalia and 
in the fore wing by the basal inner gradate 
meeting Psm. Species in the desjardinsi-group 
possess a tignum with a minute acumen, a broad 
arcessus and a cruciform, basally expanded 
gonapsis. The following species are transferred 
to Mallada and are members of the desjardinsi- 
group: anpingensis Esben-Petersen comb, n., 
meloui Navas comb. n. and maquilingi Banks ( = 
inconspicua Navas) comb. n. In a second species 
group within Mallada s. str. the gonapsis is 
absent (instead there is a large V-shaped apo- 
deme at the apex of sternite 8+9), the gonosetae 
are simple (in the desjardinsi-group the setal 
bases are thickened so that they resemble 
mounted needles), the arcessus is narrow and 
elongate and the acumen is short and ovate. It is 
now clear that some species formerly included in 
Chrysoperla belong to this second species-group 
and so are reassigned to Mallada: krakatauensis 
Tsukaguchi comb, n., satilota Banks comb. n. 
and suzukii Okamoto comb. n. 

The presence of gonocristae in australis and 
triactinata suggest a relationship with Aperto- 
chrysa, although species of this genus lack a 
tignum and possess a gonapsis. It may be neces- 
sary to erect a new genus for these two Austra- 
lian species but for the present they are 
provisionally assigned to Apertochrysa: Aperto- 
chrysa australis comb, n., Apertochrysa triacti- 
nata comb. n. 

I have been unable to examine the types of the 
following species that are deposited in BAU or 
IZAS, China: Chrysoperla quinlingensis Yang & 
Yang, C. thelephora Yang & Yang, C. yulinica 
Yang & Yang (Yang & Yang 1989), C. bellatula 
Yang & Yang, C. hainanica Yang & Yang, C. 
longicaudata Yang & Yang, C. sola Yang & 
Yang (Yang & Yang, 1992) or Chrysopa xizan- 
gana Yang (Yang, 1988). Unfortunately, it is not 
possible to decide the affinitites of these species 



from the original descriptions so at present they 
must remain incertae sedis. 

Distribution. Eight species are known from 
the Afrotropics, six from the Western Palaearctic 
region, three from the Eastern Palaearctic, there 
is one Palaearctic species, three Oriental species, 
one species occurs throughout the Old World 
tropics, there are ten Nearctic species and four 
Neotropical species. 

Key to the world Chrysoperla species 

This key does not include the following Chinese 
species which were unavailable for examination: 
Chrysoperla quinlingensis Yang & Yang, C. 
thelephora Yang & Yang, C. yulinica Yang & 
Yang, C. bellatula Yang & Yang, C. hainanica 
Yang & Yang, C. longicaudata Yang & Yang, C. 
sola Yang & Yang or Chrysopa xizangana Yang. 

1 Gradates green 2 

— Gradates black or greyish 17 

2 Base of claw undilated or only slightly swollen (Figs 
33.51) 3 

— Base of claw with distinct step-like dilation, at least 
one-quarter length of claw hook (Fig. 5) 8 

3 Head with extensive black/brown markings, with 
X-mark between antennae furcifera Okamoto 

— Markings on head restricted to red or brown stripe 
on gena. frons or clypeus 4 

4 Antennae black/brown; veins 1A and 2 A of fore 
wing black/brown mexicana sp. n. 

— Antennae green; veins 1A and 2A of fore wing 
green 5 

5 Intramedian cell (im) long, apex of im meets first Rs 
crossvein (Fig. 32) 6 

— Intramedian cell short, apex of im not reaching first 
Rs crossvein (Fig. 1) 7 

6 Post-ocular region marked red; gena marked with 
red spot; apex of fore wing pointed; cf genitalia 
with spinellae harrisii Fitch 

— Post-ocular region unmarked; gena marked with 
brown spot; apex of fore wing rounded; spinellae 
absent in c$ genitalia annae sp. n. 

7 Costal setae long, 1.5 mm; wing apex rounded (Fig. 
68); claw hook very long (Fig. 69) . renoni Lacroix 

— Costal setae short mediterranea Holzel 

8 Basal dilation of claw large, at least half length of 
claw hook 

— Basal dilation of claw small, distinctly less than half 
length of claw hook 10 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



145 



9 Pronotum marked with brown spot in each corner 
in addition to yellow median band; vertex marked 
with red lateral spot exul McLachlan 

— Pronotal markings restricted to yellow median 
band; vertex unmarked comanche Banks 

10 Gena usually marked with red spot; red post-ocular 
spot usually present; cf: spinellae present 11 

— Gena usually marked with brownish/black spot, 
which may be overlaid with red; post-ocular region 
usually unmarked; cf: spinellae absent 12 

1 1 Basal dilation of claw less than third length of claw 
hook (Fig. 189); Old World tropics cf: neck of 
acumen of tignum short, swollen apically 
congrua Walker 

— Basal dilation of claw about third length of claw 
hook (Fig. 228); Neotropics; cf: neck of acumen 
long, swollen apically externa Hagcn 

12 Fore wing long, more than 13 mm; cf: 10-20 
gonosetae lateral group annae sp. n. 

— Fore wing shorter, usually less than 13 mm; cf: few 
gonosetae (less than 10 in lateral group) 13 

13 Fore wing rounded, symmetrical apically (Fig. 40); 
Cf: membranous lip of sternite 8 + 9 narrow, elon- 
gate (Fig. 42) iranica Holzel 

— Fore wing tapered, asymmetrical apically (Fig. 3) 
(in some northern European specimens fore wing 
may be rounded with long setae Fig. 2); cf: mem- 
branous lip of sternite 8+9 short, broad, rounded 
(Fig. 8) 14 

14 Basal dilation of claw one-quarter or less length of 
claw hook (Fig. 25) 15 

— Basal dilation of claw more than third length of claw 
hook 16 

15 First Rs crossvein at oblique angle to Rs; Afrotro- 
pics zastrowi Esben-Petersen 

— First Rs crossvein usually at right angles to Rs; 
India sillemi Esben-Petersen 

— First Rs crossvein variable; Nearctic 



19 Cf: spinella present 



20 



downesi Smith 

16 Palaearctic carnea Stephens 

— Nearctic plorabunda Fitch species group 

17 Gena marked with red spot or stripe; if unmarked, 
clypeus with black spot [some specimens of the 
plorabunda species-group complex may key out at 
this point] 18 

— Gena marked brown/black, often with red overlay; 
if unmarked, clypeus with red spot 30 

18 Costal setae of fore wing long, at least 1 mm, often 
erect 19 



— Cf: spinella absent 21 

20 Gena marked with red spot, clypeus unmarked: 
pronotum marked with red stripe in each corner 
and red medio-lateral spot; claw rounded posteri- 
orly, with very large basal dilation (Fig. 134); fore 
wing less than 13 mm; cf: apex of sternite 8+9 
narrow, prominent (Fig. 135); acumen of tignum 
short, swollen apically (Fig. 136) ... plicata Tjeder 

— Gena and clypeus marked with red stripe; any red 
spots on pronotum restricted to anterior corner; 
claw straight posteriorly with smaller basal dilation 
(Fig. 125); fore wing more than 13 mm; cf: apex of 
sternite 8 + 9 short and broad (Fig. 126); acumen 
long, narrow apically (Fig. 127) .... nyerina Navas 

21 cf: arcessus striated dorsally (Fig. 79) 



oblita Holzel 



— O": arcessus without dorsal striations 22 

22 Fore wing less than 12 mm; cf: arcessus narrow 
(Fig. 104); acumen of tignum short, swollen (Fig. 
105); apex of sternite 8 + 9 prominent, longer than 
membranous lip (Fig. 101) comans Tjeder 

— Fore wing more than 12 mm; cf: arcessus broad 
(Fig. 113); acumen long, parallel-sided (Fig. 112); 
sternite 8 + 9 not elongated apically (Fig. 
Ill) decaryana Navas 

23 Maxillary palps with black dorsal stripe 24 

— Maxillary palps unmarked 27 

24 Red post-ocular spot present 26 

— Post-ocular region unmarked 25 

25 cf: apical membrane of arcessus narrow, elongate 
(Fig. 281); acumen of tignum short, broad, swollen 
apically (Fig. 280) rufdabris Burmeister 

— Cf: apical membrane of arcessus short, broad, 
tapered at apex (Fig. 216); acumen long, swollen 
apically (Fig. 215) asoralis Banks 

26 Front of vertex marked with two narrow red spots; 
Cf: arcessus abruptly widened at apex; gonarcus 
arch broad (Fig. 202) dozieri Smith 

— Front of vertex unmarked, but Cuban specimens 
may have a pair of broad red spots; cf: arcessus 
narrow apically: gonarcus arch narrow (Fig. 
22 1 ) exotera Navas 

— Front of vertex unmarked; cf: arcessus broad, 
narrowed subapically, spatulate at apex (Fig. 
206) defreitasi sp. n. 

27 Basal dilation of claw less than half length of claw 
hook 28 

— Basal dilation of claw more than half length of claw 
hook 29 



Costal setae short, inclined towards wing apex . 23 28 Basal dilation of claw about one-quarter length of 



146 

claw hook; scape with red stripe cf: spinellae 
absent gallagheri Holzel 

— Basal dilation of claw distinctly less than one- 
quarter length of claw hook; scape unmarked; cf : 
spinellae present nigrinervis sp. n. 

29 Basal dilation of claw less than two-thirds length of 
claw hook mutata McLachlan 

— Basal dilation of claw hook about two-thirds length 
of claw hook pudica Navas 

30 Gena unmarked; intramedian cell (im) long, 
extends beyond first Rs crossvein (Fig. 115) .... 31 

— Gena marked with brown spot; im short, at most Rs 
crossvein meets apex of im (Fig. 90) 32 

31 Palps marked with black dorsal stripe; black lateral 
stripe on clypeus; post-ocular region unmarked; cf : 
tignum absent; spinellae present ... insulata Fraser 

— Palps unmarked; red spot on clypeus; red post- 
ocular spot present; cf: tignum present; spinellae 
absent siamensis sp. n. 

32 Basal dilation of claw less than half length of claw 
hook (Fig. 238) ; crossveins in basal half of fore wing 
of fore wing entirely black/brown; pronotum with 
narrow black median stripe; cf: spinellae present- 

galapagoensis Banks 

— Basal dilation about half as long as claw hook; 
crossveins in basal half of fore wing black/brown at 
each end, middle section green; pronotum with 
yellow median band; cf: spinellae absent 33 

33 USA johnsoni Henry, Wells & Pupedis 

— Eastern Palaearctic nipponensis Okamoto 



SPECIES DESCRIPTIONS 

Chrysoperla carnea (Stephens) 
(Figs 1-4, 6-13, 17, 19-20) 

Chrysopa carnea Stephens, 1836: 103. Lecto- 
type?; ENGLAND: near London (BMNH) 
[examined]. [Lectotype designated by Leraut, 
1991: 78.] 

Chrysopa affinis Stephens, 1836: 104. Lecto- 
type?, ENGLAND: near London (BMNH) 
[Lectotype designation by Leraut, 1991: 78.] 
[examined]. [Synonymized by Schneider, 1851: 
68.] 

Chrysopa microcephala Brauer, 1850: 6. Holo- 
type, AUSTRIA: Vienna (NMW) [not exam- 
ined]. [Synonymized by Navas, 1908: 13.] 

Chrysopa vulgaris Schneider, 1851: 68. Syntypes, 
Europe [not traced]. [Synonymized by Killing- 
ton, 1931: 112.] 



S.J. BROOKS 

Chrysopa lamproptera Stein, 1863: 419. Syn- 
types, YUGOSLAVIA: Dalmatia, Casteln- 
uovo (Kruper) (NMHU) [not examined]. 
[Synonymized by Aspock et al., 1980: 268.] 

Chrysopa lucasina Lacroix, 1912: 203. 
Lectotypecf, ALGERIA: Frenda, 1911 
(Lucas) (MNHN) [Lectotype designated by 
Leraut, 1991: 78.] [examined]. [Synonymized 
with carnea Stephens by Aspock et al., 1980: 
268; reinstated as valid species by Leraut, 
1991: 76.] Syn. n. 

Cintameva angelnina Navas, 1931a: 84. Holo- 
type, GERMANY: Satrup (Angeln), Anfang, 
ix.1930 (Lichtwerk) (ZMH) [type destroyed]. 
[Nomen dubium; synonymised by Aspock et 
al., 1980:410.] 

Chrysopa ferganica Navas, 1933b: 107. Syntypes, 
USSR: Turkestan, Fergana, Sphara, 
25.vii.1928 (NMHU) [not examined]. [Synony- 
mized by Aspock etal., 1980: 268.] 

Chrysopa pictavica Lacroix, 1933: 147. Lecto- 
typecf; FRANCE: Deux-Sevres, marais 
d'Amure, 27. vi. 1921 {Lacroix) (MNHN) [Lec- 
totype designated by Leraut, 1982: 48.] [exam- 
ined]. [Synonymized by Aspock et al., 1980: 
268.] 

Chrysopa carnea var. adaptata Navas, 1934a: 1. 
Holotype, CHINA: Jiangsu Prov., Hufu 
(Loubou), 3.viii. 1933 (IZAS) [not examined]. 
[As subspecies of nipponensis (Okamoto) 
Yang & Yang, 1990a: 79.] Syn. n. 

Chrysopa lundbladiTjeder, 1939: 8. Holotypecf , 
MADEIRA: Caramujo, 1250m (Lundblad) 
(NRS) [not examined]. [Synonymized by 
Aspock etal., 1980:268.] 

Chrysopa maderensis Tjeder, 1939: 9. Holo- 
typecf, MADEIRA: Rabacal, 1080m, 
17.viM.viii.1935 {Lundblad) (NR) [not exam- 
ined]. [Synonymized with carnea Stephens by 
Aspock et al., 1980: 268; synonymized with 
lucasina Lacroix by Leraut, 1991: 78.] Syn. n. 

Chrysopa canariensis Tjeder, 1939: 31. Holotype 
Cf , CANARY IS: Gran Canary, Tafira {Frey) 
(NRS) [not examined]. [Synonymized by 
Aspock et al., 1980:268.] 

Chrysopa shansiensis Kuwayama, 1962: 9. Holo- 
type $, CHINA: Shansi, Wangmaochen, 
14. v. 1942 {Yasumatsu) (KU) [examined]. Syn. 
n. 

Chrysopa (Chrysoperla) carnea Stephens; Stein- 
mann, 1964: 260. 

Chrysopa (Chrysoperla) canariensis Tjeder; 
Tjeder, 1966: 391. 

Chrysopa (Chrysoperla) lundbladi Tjeder; 
Tjeder, 1966: 391. 

Anisochrysa (Chrysoperla) carnea (Stephens); 
Holzel, 1970: 51. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



147 



Chrysoperla carnea (Stephens); Canard & Lau- 
deho, 1978:71. 

Chrysopa carnea nanceiensis Semeria, 1980: 29. 
Syntypes, FRANCE: Nancy (Poivre) 
(NANCY UNIV.) [not examined]. [Synony- 
mized by Leraut, 1991: 80; reinstated as valid 
species by Semeria, 1992: 337.] Syn. n. 

Chrysoperla shansiensis (Kuwayama); Brooks & 
Barnard, 1990: 271. 

Fore wing (Fig. 1) d\ 9-13 mm; 9- 11-14 mm. 
Head marked with brown/black stripe on gena 
and lateral clypeus, often overlaid with red 
stripe; post-ocular spot absent. Maxillary palps 
unmarked or with black stripe dorsally. Anten- 
nae shorter than fore wing. Pronotum marked 
with yellow/white longitudinal median stripe, 
greyish spot sometimes present in anterior cor- 
ner; prothoracic setae short, dark or sometimes 
pale. Claw (Fig. 4) with small basal dilation, 
about one-third length of claw hook (2.86-3.0: 1 ; 
mean 2.70). Fore wing variable in width (length: 
breadth = 2.7-3.1: 1); pointed or rounded api- 
cally; venation green but costal and Rs crossveins 
often with black spot at Sc and R respectively 
and crossveins im-Psm, cl-c2, Rs-im sometimes 
entirely black; costal setae short or relatively 
long (0.4-1.6 mm), inclined towards wing apex; 
cell im short, not reaching 1st Rs crossvein or 
sometimes meeting at apex; 1st Rs crossvein 
meets Rs obliquely or at 90°; 4-7 inner gradates, 
5-8 outer gradates. 

Genitaliao" (Figs 8-10). Lip of sternite 8 + 9 
broad, rounded. Apodeme of tergite 9 long, 
extending well-beyond callus cercus; callus cer- 
cus broad, ovate; 23-33 trichobothria. Spinellae 
absent. Gonosetae few, 4-6 in lateral clump, 
often asymmetrical in number. Arcessus striated 
dorsally; tip blunt; apex straight or curved ven- 
trally at 45-90°. Entoprocessus large, crescentic. 
Gonarcus with large, reniform lateral plates; 
arch broad. Acumen of tignum long, rounded, 
not swollen, apically. 

GENITALIA9 (Figs 11-13). Callus cercus ovate; 
37 trichobothria. Subgenitale with short basal 
extension. Spermatheca with deep ventral 
impression; vela long; duct long. 

Remarks. The male genitalia of carnea are 
practically indistinguishable from zastrowi, medi- 
terranean furcifera, oblita, nipponensis and 
plorabunda and these species are obviously 
closely related to each other. However, carnea 
can usually be differentiated from these species 
by the combination of green gradate crossveins 
and relatively broad wings, small basal dilation of 



the claws and short, black prothoracic setae. 

Chrysopa shansiensis Kuwayama is treated as a 
synonym of carnea for the first time in this work. 
Unfortunately, all that remains of the holotype 
of shansiensis is the extreme apex of the abdo- 
men (sternite 8, the ectoprocts and spermatheca) 
and from this it is impossible to be certain of the 
true identity of the species. In his description of 
shansiensis, Kuwayama (1962) compares the spe- 
cies with Nineta flava (Scopoli) and N. vittata 
(Wesmael). However, the wing venation shown 
in the photograph of the complete holotype of 
shansiensis (Kuwayama, 1962) does not resemble 
species of Nineta. Rather, the specimen appears 
to be a Chrysoperla species and the description 
suggests that it is a weakly marked example of 
carnea. 

Yang & Yang (1990a) considered C. carnea 
var. adaptata Navas to be a subspecies of nippon- 
ensis. They based their decision on similarities in 
the female genitalia between nipponensis and 
adaptata without indicating what those similari- 
ties were. I have found the female genitalia of 
nipponensis and carnea to be indistinguishable. 
One character that readily serves to distinguish 
carnea and nipponensis. however, is that the 
gradate crossveins are green in carnea but black 
in nipponensis. The gradates of adaptata are 
green so it seems logical to treat this taxon as a 
synonym of carnea. 

Geographical variation. Mediterranean. A 
series of specimens examined from Crete, Sar- 
dinia, central Greece (Amfissa. Fokidhos). the 
Peloponnese and Spain have distinctive mark- 
ings. The most striking features, which distin- 
guish this population from typical carnea, are the 
very pale green overall ground colour of the body 
and a pair of conspicuous black dorso-lateral 
spots on the basal segments of the abdomen. In 
addition, the head lacks any red markings and 
the longitudinal median stripe on the pronotum 
and abdomen is white. All crossveins are black at 
each end, rather than being entirely green, the 
wings may be pointed or rounded apically with 
short or long costal setae respectively. The basal 
dilation of the claw is larger than typical for 
carnea (Fig. 6), being only slightly under half the 
length of the claw hook (2.06-2.64: 1: mean 
2.42). 

Israel. A single male specimen from Lake 
Huleh on the River Jordan was examined that 
differed in several respects from typical carnea, 
although the genitalia were indistinguishable 
from typical carnea. Head suffused entirely red; 
gena marked with a faint brown stripe; claw with 
large basal dilation (Fig. 7) (2.41: 1); costal setae 



148 



S.J. BROOKS 



long, semi-erect; fore wing apex rounded. How- 
ever, the lack of further material precludes the 
description of this specimen as a distinct taxon. 

Europe. Leraut (1991; 1992) drew attention to 
the co-occurrence of three forms of carnea in 
Europe. The first, which Leraut established as 
carnea s.str., has a dark brown to black stripe on 
the gena, rounded apex to the fore wing (Fig. 2) 
and relatively long wing setae. This form is 
dominant in northern European populations but 
occurs throughout the distributional range. A 
second form was referred to as carnea B (Leraut, 
1991) but was later elevated to species, and 
identified as C. kolthoffi (Navas) (Leraut, 1992). 
This taxon was said to differ from carnea s. str. in 
having short, dark setae on the wing veins and 
narrow arms in the tignum of the male genitalia. 
However, I was unable to detect any difference 
between specimens identified by Leraut as 
kolthoffi and those identified as carnea. The 
narrow arms of the tignum shown by Leraut 
(1991) could be due to individual variation or 
distortion, as is often the case in teneral or 
immature specimens. In any case, C. kolthoffi is 
an incorrect name for this taxon since kolthoffi is 
a synonym of the Eastern Palaearctic species, 
Chrysoperla nipponensis (Okamoto). The third 
form, which Leraut (1991) referred to as C. 
lucasina (Lacroix), differs from C. carnea in 
having a paler stripe on the gena, short costal 
setae and a straighter apico-posterior margin in 
the fore wing which makes the wing appear more 
pointed (Fig. 3). This form is dominant in south- 
ern Europe, north Africa and the Middle East. 

However, the morphological differences indi- 
cated by Leraut are not as clear-cut as he sug- 
gests. First, I have found every stage of genal 
marking from red to dark brown on specimens 
that would otherwise be referrable to lucasina 
and, conversely, carnea-type specimens with a 
pale brown stripe on the gena. Secondly, I have 
examined several different contemporaneous 
and sympatric series from various European 
localities that show a gradation from the pointed 
wing form with short setae to the rounded wing 
form with long setae. Other specimens were 
examined in which the fore wing was rounded 
but costal setae short and, conversely, specimens 
in which the fore wing was pointed apically and 
the costal setae long. In addition, I examined 
several specimens that exhibited an intermediate 
condition and it was not possible to decide 
whether the wings were pointed or rounded 
apically. 

In North America there is similar variation in 
specimens of C. plorabunda with some speci- 
mens having rounded wing apices with long cos- 



tal setae but the majority having more pointed 
wings and short costal setae. No distinction has 
yet been found in the courtship songs of round 
wing and pointed wing forms of C. plorabunda. 

No significant difference was found in the size 
of the basal dilation of the claw in specimens 
referrable to f. typica or f. lucasina. 

At present, I do not consider the morphologi- 
cal evidence conclusive enough to treat C. luca- 
sina and C. carnea as distinct species. However, 
further taxonomic and biological investigations 
will help to clarify this problem. 

As an interesting addition to these consider- 
ations, Barnard et al. (1986) reported the sympa- 
tric occurrence of two forms of carnea larvae, 
which had different head markings, on oak trees 
in southeast England. However, although a long 
series of adults was also collected throughout the 
same year they were all f. typica with rounded 
wing apices and long wing setae. 

Using a multivariate statistical method Thierry 
et al. (1992) discriminated three morphs of car- 
nea in populations in central France which they 
referred to as carnea, lucasina and kolthoffi. The 
three main variables used to produce their results 
were the colour and length of the costal setae on 
the fore wing, the colour and distribution of 
prothoracic setae, and the shape of the spot on 
the stipe. These interesting results point the way 
to further research that may establish the pres- 
ence of sibling species among populations of 
carnea but at present it is premature to recognise 
these taxa as valid species. It is beyond the scope 
of this study to conduct extensive multivariate 
analysis of Chrysoperla species. 

Semeria (1992) carried out a morphometric 
analysis of the male genitalia of some French 
specimens in the carnea species group. Essen- 
tially he examined the size of the lateral plates of 
the gonarcus. This was done by measuring the 
angles between lines drawn from the apex of the 
arcessus to the apex of the lateral plates (BAC) 
and the angle between lines from the apex of the 
lateral plates to the mid-point of line drawn along 
the base of the lateral plates (BDC). Semeria 
considered that the results indicated the presence 
of six distinct species (C. renoni, lucasina, medi- 
terranea, carnea, nanciensis and carnea sp. B) in 
France. However, I carried out a similar analysis 
of carnea group specimens and produced results 
that differed widely from those of Semeria 
(1992). 

Because Semeria does not record any variation 
in the angles he measured I assume that the 
results he obtained for each taxon were derived 
from only one specimen. My results show consid- 
erable variation between specimens and overlap 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



149 



the range measurements given by Semeria for all 
the taxa he investigated. I consider it highly 
unlikely that each one of the specimens I exam- 
ined is a distinct species in its own right and is 
also different to the taxa investigated by Seme- 
ria. While the technique advocated by Semeria is 
worthy of further investigation it is desirable that 
a large number of specimens are examined 
before any taxonomic decisions can be based on 
the results. Consideration should also be given to 
factors that my affect the results such as the 
effects of compressing the genitalia on a micro- 
scope slide. The maturity of the specimen may 
also effect the results. Adams & Penny (1987) 
have suggested that the genitalia may become 
more heavily sclerotised as the insect matures. 
This implies that the lateral plates of the gonar- 
cus may actually increase in size as an individual 
ages. 

The morphology of the tignum shows consider- 
able variation throughout the range of C. carnea 
(Figs 17-20) both in length of the arms and 
length of the acumen. Although I could not 
detect any obvious pattern in this variation this 
may merit further investigation. 

Henry (1985a) reports the occurrence of two 
distinct song morphs of carnea in Europe. One, 
'Basel carnea\ from lowland meadows and oak 
scrub in France, Switzerland and south to 
Morocco. A second morph, "Ticino carnea', 
occurs in the southern Swiss Alps. These taxa 
cannot be distinguished from one another mor- 
phologically. 

Distribution. Palaearctic. 

Material examined (429d\ 6959) Europe: 
Belgium, Corsica, Crete, Cyprus, Czechoslova- 
kia, Denmark, France, Germany, Italy, Norway, 
Switzerland, Portugal, Sardinia, Spain, UK, 
Yugoslavia; North Africa: Algeria, Canary Is, 
Egypt, Libya, Madeira, Tunisia; Middle East: 
Iran. Iraq, Jordan, Saudi Arabia, Turkestan, 
Turkey, Yemen; India (Kashmir); China. 

Ohm & Holzel (1992) record C. carnea from 
Sudan and Senegal but all specimens examined in 
this study from the Sahel region of Africa were 
referrable to C. zastrowi. 

Chrysoperla sillemi (Esben-Petersen) stat. 
rev. 

(Figs 5, 8) 

Chrysopa sillemi Esben-Petersen, 1935: 234. 
Holotype °- INDIA: Kashmir, Karakorum, 
Maralbashi, 110m, 20.iii.1930 (Sillem) (ITZ) 
[examined]. Stat. rev. 



Chrysopa (Chrysoperla) gujaratensis Ghosh, 
1976a: 74. Holotype o\ INDIA: Gujarat, 
Dwarka, E. of Rly Stn, 2.xi.l973 (Gupta) 
(ZSI) [not examined]. Syn. n. 

Chrysopa (Chrysoperla) punensis Ghosh, 1976b: 
189. Holotype d\ INDIA: Maharashtra, 
Khed, R. Bhama, Pune, 3.vii.l976 (Ghosh) 
(ZSI) [not examined, paratype examined]. 
Syn. n. 

Chrysopa (Chrysoperla) sanandensis Ghosh, 
1977: 103. Holotype d". INDIA: Gujarat, 
Dadagram, Sanand, 8.xii. 1973 (Cherian) (ZSI) 
[not examined, paratype examined]. Syn. n. 

Fore wing 0", 10-13 mm; 9- U-14 mm. Head 
occasionally unmarked or with pale brown stripe 
on gena and clypeus, overlaid with red stripe. 
Maxillary palps brown dorsally. Antennae 
shorter than fore wing. Pronotum with yellow or 
white median stripe; setae short (0.6 mm), dark 
or pale. Claw (Fig. 5) with small basal dilation 
3.60-5.13: 1 (mean 4.17). Fore wing length; 
breadth = 2.7-3.3: 1; venation green; apex 
pointed; costal setae short, inclined towards wing 
apex. 

GENITALIA d* and 9 as Ior C. carnea except in 
male genitalia, acumen of tignum short (Fig. 8). 

REMARKS. C. sillemi can be reliably distin- 
guished from C. carnea only by differences in the 
size of the basal dilation of the claw. In C. sillemi 
the basal dilation is about one quarter the length 
of the claw tooth but in C. carnea the basal 
dilation is at least one third the length of the claw 
tooth. 

Because the ZSI has a policy of not lending 
holotypes I was unable to borrow the types of the 
Chrysoperla species described by Ghosh. How- 
ever, paratypes were available for C. punensis 
and sanandensis. These specimens were typical 
examples of C. sillemi. Unfortunately, no 
paratypes were available for C. gujaratensis. 
Nevertheless, from the good description and 
figures provided for this species (Ghosh, 1976a) 
there can be little doubt that it is synonymous 
with C. sillemi. The setae shown on the tignum 
probably originate from the subanale. 

Distribution. India, Pakistan. Specimens of 
both C. sillemi and C. carnea were examined 
from Kashmir. 

Material examined (23d", 309) India, Paki- 
stan. 



150 



S.J. BROOKS 



Chrysoperla plorabunda (Fitch) 

(Figs 15, 21-22) 

Chrysopa plorabunda Fitch, 1855: 792. Syntypes, 
USA: New York State & Illinois (MCZ) [not 
examined]. [Synonymized with carnea 
Stephens by Banks, 1903: 155; reinstated as 
valid species by Henry, 1983: 299.] 

Chrysopa robertsonii Fitch, 1855: 792. Syntypes, 
USA:Arkansas, Tullehassie (MCZ) [not 
examined]. [Synonymized by Banks, 1903: 
162.] 

Chrysopa pseudographa Fitch, 1855: 793. Syn- 
types, USA: Illinois (MCZ) [not examined]. 
[Synonymized by Banks, 1903: 162.] 

Chrysopa dlinoiensis Shimer, 1865: 208. USA: 
Illinois[not examined]. [Synonymized by 
Banks, 1903: 155.] 

Chrysopa californica Coquillet, 1890: 288. Holo- 
type, USA [holotype not traced]. [Smith, 1932: 
594 as synonym of plorabunda; Adams, 1956: 
45 as subspecies of plorabunda.] Syn. n. 

Remarks. Morphologically Chrysoperla 

plorabunda is indistinguishable from C. carnea. 
However, Henry (1983; 1985a; 1985b and in litt.) 
has provided evidence that these two taxa should 
be regarded as distinct species. The substrate- 
borne songs of European populations of carnea 
are completely different from those of eastern 
and western New World populations of 
plorabunda. Egg production resulting from 
crosses of the Old and New World populations of 
these two taxa was very low. Although Henry's 
data is derived from relatively few populations it 
is persuasive and I am reluctant to synonymise 
the species solely on morphological grounds. The 
accrual of more song data from European and 
American populations will clarify the issue. 

Like carnea, specimens of plorabunda varied 
in the length of the arms and acumen of the 
tignum. In addition, the number of gonosetae 
varied from four to eight in each lateral group. 
This is a greater range of variation than that 
noted in carnea. The basal dilation of the claw of 
C. plorabunda is similar in size to that of C. 
carnea (2.4-3.09: 1; mean 2.83) 

Distribution. Nearctic. 

Material examined (22cf, 38$) Canada, 
Mexico, USA (Arizona, California, N. & S. 
Carolina, Colorado, Florida, Montana, 
Nebraska, Oregon, Texas, Utah, Vermont, 
Washington, Wisconsin). 



Chrysoperla johnsoni Henry, Wells & 
Pupedis 

(Fig 23) 

Chrysoperla johnsoni Henry et al., 1993:10. 
Holotype d\ USA: Oregon, Philomath 
(Mary's Peak), 28.ix.1989 (Henry & Wells) 
[not examined]. 

Fore wing d\ 10-11.5 mm; $, 11-13.5 mm. 
Head marked with red stripe on gena, sometimes 
with additional brown stripe; red lateral stripe on 
frons and clypeus; red post-ocular spot present or 
absent. Maxillary palps marked brown dorsally. 
Antenna shorter than fore wing. Pronotum with 
yellow median longitudinal band and greyish 
lateral stripe, sometimes with reddish brown 
mid-lateral spot; prothoracic setae short, pale or 
dark. Claw (Fig. 15) with large basal dilation, 
about half length of claw hook (2.10-2.82: 1: 
mean 2.34). Fore wing narrow or broad (length : 
breadth = 2.9-3.4 : 1), rounded apically; costal 
setae short (0.6-0.8 mm), inclined towards wing 
apex; 1st Rs crossvein meets Rs obliquely; cell im 
short, not meeting 1st Rs crossvein; 4—6 inner 
gradates, 5-7 outer gradates; gradates green or 
black; crossveins green or black at each end 
(including at wing margins); R marked with short 
black stripe basal of tympanal organ. Abdomen 
unmarked or marked with brown dorso-lateral 
spot on segment two. 

Genitalia d" and 9 indistinguishable from 
Chrysoperla carnea. 

REMARKS Chrysoperla johnsoni was distin- 
guished from C. adamsi and C. plorabunda on 
the basis of differences in their courtship songs 
(Henry et al., 1993; Henry, 1993). There are two 
morphs of C. johnsoni. The typical form, which 
is indistinguishable morphologically from C. 
plorabunda, has green gradates and the basal 
dilation of the claw is about 2.8:1. The second 
form, at present known only from the San Fran- 
cisco Bay area, has black gradate crossveins, the 
basal dilation of the claw is usually broad (about 
2.2:1), the head has extensive red markings and 
the gena often lacks a brown stripe. This form is 
morphologically indistinguishable from the 
"mohave' morph of C. downesi. 

Distribution. Western USA 

Material examined (7o\ 15$) USA: Ari- 
zona, California. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



151 



Chrysoperla adamsi Henry, Wells & 
Pupedis 

Chrysoperla adamsi Henry et al., 1993: 9. Holo- 
type, 0\ USA: Oregon, Philomath (Mary's 
Peak), 28. ix. 1989 (Henry & Wells) [not exam- 
ined]. 

Fore wing 12-13 mm. Head marked with reddish 
brown spot on gena; red stripe on gena, lateral 
frons, lateral clypeus. Maxillary palps marked 
black dorsally. Pronotum and vertex of head with 
yellowish white median stripe; setae short, black. 
Basal dilation of claws less than half length of 
claw hook (2.54: 1). Wing venation green; fore 
wing relatively narrow (length: breadth = 
2.9-3.2: 1); apex pointed; costal setae short 
(0.4-0.6 mm), inclined towards wing apex; im 
short, not reaching 1st Rs crossvein; 6 inner 
gradates, 6-7 outer gradates. 

Genitalia o\ 9 as described for C. carnea. 

Remarks. C. adamsi cannot be separated mor- 
phologically from C. plorabunda, C. johnsoni or 
C. downesi. However, Henry et al. (1993) has 
distinguished this taxon on the basis of its court- 
ship song. Unlike C. johnsoni, which may have 
black gradate crossveins, specimens of C. adamsi 
have green wing venation. 

Distribution. Western USA 

Material examined (lo\ 1$) USA: Califor- 
nia [material identified by C.S. Henry]. 

Chrysoperla downesi (Smith) stat. rev. 

(Figs 14, 16) 

Chrysopa downesi Smith, 1932: 594. Holotype 9 
CANADA: British Colombia, Kelowna 
(CNC) [not examined]. [Synonymized with 
carnea Stephens by Garland, 1985a: 93.] Stat, 
rev. 

Chrysopa mohave Banks, 1938: 120. Holotype, 
USA (MCZ)[not available for examination]. 
[Synonymized with C. carnea by Tauber & 
Tauber, 1973a: 1164.] Syn. n. 

Fore wing (Fig. 14) d\ 11.5 mm; $, 
12-12.5 mm. Head marked with black stripe on 
gena; red lateral stripe on frons and clypeus; red 
post-ocular spot often present. Maxillary palps 
black. Antenna shorter than fore wing. Prono- 
tum marked with white longitudinal stripe, lat- 
eral margins sometimes marked reddish brown; 
prothoracic setae short, black. Claw (Fig. 16) 
with basal dilation usually less than one-third 
length of claw hook (3.7-5.17: 1; mean 4.26) but 



specimens with black gradates have a large basal 
dilation (2.4-2.7:1). Fore wing narrow (length: 
breadth = 3.0-3.5: 1): pointed apically; venation 
mostly green but c,lc 2 crossvein black, gradates 
green or black; costal setae short (0.4-0.8 mm), 
inclined towards wing apex: cell im short, not 
reaching 1st Rs crossvein or sometimes meeting 
apex of cell: 1st Rs crossvein meets Rs obliquely 
or at 90°; 4-6 inner gradates, 5-7 outer gradates. 

Genitalia, c/ and 9 and genitalia indistin- 
guishable from C. carnea. 

Remarks C. downesi and C. plorabunda are 
very similar morphologically and it is not always 
possible to distinguish the two species. However, 
some specimens of downesi, especially those 
from the western USA, are marked with a red 
post-ocular spot between the eye and vertex. 
This marking is never present in plorabunda. 
Other characters which serve to distinguish the 
two species are the white longitudinal protho- 
racic band in downesi. which is yellow in 
plorabunda, the basal dilation of the tarsal claw, 
which is generally smaller in downesi than 
plorabunda, and the width of the fore wing which 
tends to be narrower in downesi than 
plorabunda. However, these characters are 
prone to variation and cannot be relied upon to 
separate the species. Earlier workers (e.g. 
Tauber & Tauber. 1981) have considered C. 
downesi to have a dark green ground colour but, 
following courtship song analysis, Henry (1992) 
has shown that some populations of C. downesi 
include yellow individuals. 

Tauber & Tauber (1977a) and Henry (1985a; 
1985b) recognised downesi as a valid species on 
the basis of ecological evidence. C. downesi 
occurs in pine woodland and has a univoltine life 
cycle with a summer diapause. In life, the species 
has a year-round dark-green ground colour with 
a whitish median longitudinal band and is often 
covered with an oil-like secretion that is detect- 
able in museum specimens. C. plorabunda occurs 
in deciduous woodland and open meadow habi- 
tats and has a multi-voltine life cycle with no 
summer dipause. The species is pale green in the 
summer but during the winter, C. plorabunda 
becomes reddish brown. Tauber & Tauber 
(1977a; 1977b) have postulated that downesi 
evolved from plorabunda through a process of 
habitat divergence and seasonal isolation. Henry 
(1985b) showed that the courtship songs of 
downesi and plorabunda were sufficiently differ- 
ent to cause reproductive isolation between the 
two taxa in the field. However, Henry (1985b) 
and Tauber & Tauber (1982a) were able to 



152 



S.J. BROOKS 



hybridise downesi and plorabunda in the labora- 
tory, although acceptance rates of heterospecifics 
were at low levels (10-30%). Hybridisation does 
not appear to occur in the wild (Henry, 1992). 

Henry (1985a) reports on the occurrence of 
three distinct song morphs of downesi in the 
mountains of western USA. Morph Dl is wide- 
spread from eastern Canada to western Montana 
and south to California. However, this morph 
occurs sympatrically with a morphologically 
indistinguishable song morph (D2) in the north- 
ern Rocky Mountains. D2 co-occurs with a third 
song morph (D3) in central California which also 
cannot be separated using morphological charac- 
ters. 

Garland (1985b) states that the female speci- 
men bearing the holotype label has a data label 
indicating that it was collected in Vernon rather 
than in Kelowna as stated in the original descrip- 
tion (Smith, 1932). 

The true status of C. mohave has been much 
discussed in the literature (Tauber & Tauber, 
1973a; Henry, 1993). Morphologically the taxon 
differs from C. plorabunda in having the cross- 
veins black at each end and the gradates com- 
pletely black. Tauber & Tauber (1973b; 1982b; 
1986; 1987) recognised mohave as a distinct 
'strain' of C. plorabunda, with which it hybri- 
dises, on ecophysiological and life history 
grounds. Henry (1992; 1993) investigated the 
courtship song of specimens referrable to C. 
mohave and found that they fell into two distinct 
groups. One group was indistinguishable from C. 
johnsoni, the other was the same as C. downesi. 
From a taxonomic point of view it is now impos- 
sible to decide whether C. mohave should be 
considered synonymous with C. johnsoni or C. 
downesi. In order to preserve nomenclatural 
stability I have decided to consider C. mohave as 
a synonym of C. downesi. 

Distribution. Nearctic. 

Material examined (31d\ 18$) USA: Cali- 
fornia, Colorado, New York, Oregon, Texas, 
Utah, Vermont, Wisconsin. 



Chrysoperla zastrowi (Esben-Petersen); Holzel, 
1989b: 167. 

Fore wing (Fig. 24) c? 10.5-13 mm; $ 
9.5-13 mm. Head sometimes unmarked or 
marked with dark brown (occasionally reddish) 
spot on gena; reddish brown lateral stripe on 
frons and clypeus; maxillary palps brown. Anten- 
nae shorter than fore wing. Pronotum with 
median longitudinal yellow band; sometimes 
with red or brown blotches in anterior corner. 
Basal dilation of claw small (4.0-5.0: 1; mean 
4.38) Wing venation green; fore wing broad 
(length: breadth = 2.8-3.1: 1); costal setae mod- 
erately long (0.6-1.2 mm); 1st Rs crossvein 
leaves Rs at oblique angle. 

Genitaliao" (Figs 26-28). Callus cercus 
rounded; 27-37 trichobothria. Gonosaccus with 
minute spicules; few gonosetae, about 5-6 
arranged in lateral clump; apical membrane of 
arcessus short, blunt abruptly curved ventrad. 

Genitalia? (Figs 29-31). Callus cercus round; 
32 trichobothria. Subgenitale with short basal 
extension. Spermatheca wih deep ventral impres- 
sion; vela long; duct long. 

REMARKS. C. zastrowi is very similar to C. cor- 
nea but may be distinguished by the basal dila- 
tion of the claw, which is smaller in C. zastrowi 
than C. carnea. Tjeder (1966) notes that in C. 
zastrowi the 1st Rs crossvein is obliquely angled 
whereas in carnea this crossvein is at right-angles 
to Rs. However, this character is not always 
reliable since several Palaearctic specimens oth- 
erwise referrable to carnea have an oblique cross- 
vein. Nevertheless, in all southern African 
specimens examined the 1st Rs crossvein was 
oblique and never at right-angles to Rs. The male 
genitalia of C. zastrowi and C. carnea are indis- 
tinguishable. 

Distribution. Afrotropical. 

Material examined (17d\ 13$) . Ascension 
I., Namibia, Somalia, St Helena, South Africa, 
Sudan, Zambia. 



Chrysoperla zastrowi (Esben-Petersen) 

(Figs 24-31) 

Chrysopa zastrowi Esben-Petersen, 1928a 
220.Holotype $ [destroyed], NAMIBIA 
Grootfontein (ZMH). Neotype $ NAMIBIA 
Orupembe (LUND) [designated by Tjeder 
1966: 403.] [not examined]. 

Chrysopa (Chrysoperla) zastrowi (Esben 
Petersen); Tjeder, 1966: 402. 



Chrysoperla furcif era (Okamoto) 

(Figs 32-39) 

Chrysopa furcifera Okamoto, 1914: 61. Lecto- 
type $, TAIWAN: Shirin, 31.vii.1907 (Mat- 
sumura) (HU) [examined]. [Lectotype 
designated by Kuwayama, 1966: 136.] 

Chrysopa savioi Navas, 1933c: 4. Holotype $, 
CHINA: Kowloon (IZAS) [not examined]. 
Syn. n. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



153 



Chrysopa kulingensis Navas, 1936b: 56. LECTO- 
TYPE 0" HERE DESIGNATED. CHINA: 
Ruling (IZAS) [examined]. [Synonymized 
with savioi by Yang et al., 1990a: 82.]. Syn. n. 

Chrysopa (Chrysoperla) furcifera Okamoto; 
Tjeder, 1966: 391. 

Chrvsoperla furcifera (Okamoto); Tsukaguchi, 
1985: 504. 

Anisochrysa (Chrysoperla) furcifera (Okamoto); 
Holzel, 1970:51. 

Chrysoperla savioi (Navas); Brooks & Barnard, 
1990:271. 

Fore wing (Fig. 32) cf, 11.5-12.5 mm; $, 
15 mm. Head with broad black lateral stripe 
across gena, clypeus, labrum; in some specimens 
clypeus entirely black; large, black or reddish 
brown X-marking between antennae with arms 
extending onto lateral margins of vertex and 
epistomal suture; scape marked with black stripe 
on inner and outer lateral surface; maxillary 
palps black. Antennae shorter than fore wing; 
flagellar segments broad (length: breadth = 2: 
1). Pronotum with broad brown lateral band and 
yellow longitudinal median band; prothoracic 
setae short, black. Claw (Fig. 33) undilated 
basally. Fore wing narrow (length: breadth = 
3.4-3.7: 1), pointed at apex; venation entirely 
green; costal setae short or long (0.6-1.6 mm), 
erect or inclined towards wing apex; cell im 
relatively broad, 1st Rs crossvein meets im at 
apex or subapically; 4-10 inner gradates, 9-10 
outer gradates; basal inner gradates not meeting 
Psm. Abdominal setae coarse, dense. 

Genitalias (Figs 34-36). Dorsal apodeme of 
tergite 9 long, extending well beyond callus cer- 
cus; callus cercus broadly ovate or circular; 24-36 
trichobothria. Lip of sternite 8+9 broad, 
rounded. Spinellae absent. Gonosaccus with 
minute spicules; few gonosetae, 6-15 in central 
clump; arcessus striated dorsally; apex tapering 
to point, abruptly curved ventrally. Entoproces- 
sus large, crescentic. Gonarcus with large boot- 
shaped lateral plate; arch broad. Acumen of 
tignum long, narrow, slightly swollen apically. 

Genitalia? (Figs 37-39). Callus cercus round; 
36 trichobothria. Subgenitale straight basally. 
Spermatheca with shallow ventral impression; 
duct long; vela long. 

Remarks. C. furcifera is one of the most distinc- 
tive species in the genus and can be readily 
recognized by the extensive black head markings 
and narrow, pointed fore wings. Other distin- 
guishing characters include the lack of a basal 
dilation of the claws, broad flagellar segments. 



dense abdominal setae and the inner basal gra- 
date not meeting the pseudomedian vein. The 
lack of spinellae, striated arcessus, large entopro- 
cessus and relatively few gonosetae indicate that 
furcifera is related to the carnea-group of species. 
Yang & Yang (1992) distinguished the Chinese 
species savioi from the Japanese furcifera on the 
basis that the clypeus is marked black in savioi 
but unmarked in furcifera. However, in one 
Chinese example (of nine specimens examined in 
this study) the clypeus was only faintly brown 
(possibly an immature specimen) and in one 
Japanese specimen the clypeus was entirely red- 
dish brown. No differences were detected in the 
genitalia of these two populations. In view of this 
the differences in markings noted by Yang & 
Yang (1992) do not seem to me to be sufficient to 
regard savioi and furcifera as distinct species. 

Geographic variation Specimens examined 
from China and Hong Kong differed from other 
material in having darker, more extensive head 
markings. In these specimens the head markings 
are usually black rather than reddish brown. In 
addition, the inner stripe on the scape is consid- 
erably broader than the outer stripe and the 
clypeus is usually (but not invariably) entirely 
black. In specimens examined from other regions 
the clypeus is unmarked or suffused with red. 
Similarly, the Chinese specimens had dark brown 
antennae whereas material from other regions 
had the antennae pale brown. In addition, it was 
noted that the fringe of costal setae was generally 
longer in specimens from China and the Philip- 
pines (1.2-1.6 mm) than in specimens from 
Hong Kong and Japan (0.6-1.0 mm). 

Distribution. Bonin I. (Adams, 1959), China, 
Hong Kong, Japan, Philippines, Taiwan. 

Material examined (6cf , 10$) . China, Hong 
Kong, Japan, Philippines, Taiwan. 

Chrysoperla iranica (Holzel) 
(Figs 40-49) 

Chrysopa (Chrysoperla) iranica Holzel, 1967: 33. 
Holotype cf, IRAN: Vanak, 1600m, 15km 
north of Teheran, l-10.vii.1962 (Vartian) 
(HOLZEL COLL) [examined]. 

Anisochrysa (Chrysoperla) iranica (Holzel); Hol- 
zel, 1970:51. 

Chrysoperla iranica (Holzel); Brooks & Barnard, 
1990: 271. 

Fore wing (Fig. 40) d\ 9.5-10 mm; $, 10.5 mm. 
Head marked with narrow, dark brown stripe on 
gena and lateral clypeus. Maxillary palps black 



154 



S.J. BROOKS 



dorsally. Antennae shorter than fore wing. 
Pronotum marked with median longitudinal 
white/yellow stripe; prothoracic setae short, pale. 
Claw (Fig. 41) with small basal dilation, about 
one-third length of claw hook (2.72: 1). Fore 
wing broad (length: breadth = 2.7-3.1: 1), 
rounded apically; costal setae quite long 
(0.8-1.0 mm), inclined towards wing apex; 4-6 
inner gradates, 5-7 outer gradates; gradates 
green; crossveins black at each end, except costal 
and Psm crossveins which are green at the wing 
margins; anal veins green; cell im variable in 
length. Abdominal tergites with white/yellow 
median longitudinal band. 

Genitalias (Figs 42-46). Lip of sternite 8+9 
elongate, truncate apically with subapical con- 
striction. Apodeme of tergite 9 long, extends far 
beyond callus cercus; callus cercus narrow, 
ovate; 27 trichobothria. Spinellae absent. 
Gonosetae few, 5-9 in lateral clump. Arcessus 
with dorsal striations; apex blunt; apical mem- 
brane short, bent ventrad at 90°. Entoprocessus 
large, crescentic. Gonarcus with lateral plate 
broad, reniform; arch broad. Acumen of tignum 
narrow, rounded apically. 

Genitalia? (Figs 47-49). Callus cercus ovate; 
30 trichobothria. Subgenitale with narrow, short 
basal extension. Spermatheca with shallow ven- 
tral impression; duct long; vela long. 

Remarks. C. iranica belongs to the carnea- 
group of species. However, iranica can be distin- 
guished from these species by the broad fore 
wing with rounded apex and, in males, by the 
extraordinary elongation of the lip on sternite 
8+9. 

Distribution. Iran. 

Material examined (3cf , 3$) Iran. 

Chrysoperla mediterranea (Hdlzel) 

(Figs 50-58) 

Anisochrysa (Chrysoperla) mediterranea H61- 
zel, 1972: 81. Holotype cf, TUNISIA: Biserta, 
100m, 9.viii.l971 (Gepp) (HOLZEL COLL.) 
[not examined]. 

Anisochrysa mediterranea (Holzel); Monser- 
rat, 1977: 130. 

Chrysoperla mediterranea (Holzel); Aspock et 
al., 1980: 269. 

Fore wing (Fig. 50) cf , 8.5-9.5 mm; $, 11 mm. 
Head sometimes entirely suffused red or marked 
with red/brown stripe on gena, frons and clypeus; 
labrum sometimes red; red post-ocular spot 
sometimes present. Maxillary palps black dor- 



sally or unmarked. Antennae shorter than fore 
wing. Pronotum marked with longitudinal 
median yellow stripe and sometimes with brown 
median dorso-lateral spot; prothoracic setae 
short, black or pale. Claw (Fig. 51) with minute 
basal dilation (8.25: 1). Fore wing narrow 
(length: breadth = 3.2^4.0: 1); venation green; 
costal setae short (0.4-0.6 mm), inclined towards 
wing apex; cell im short, apex of cell not reaching 
1st Rs crossvein; 4-6 inner gradates, 5-7 outer 
gradates. 

Genitaliao" (Figs 52-55). Membranous lip of 
sternite 8+9 broad, rounded. Apodeme of terg- 
ite 9 long, extending well beyond callus cercus; 
23-24 trichobothria. Spinellae absent. Gonosetae 
few, 5-6 in lateral clump. Arcessus with dorsal 
striations; straight but extreme apex bent ventrad 
at 90°, apex blunt. Entoprocessus large, crescen- 
tic. Lateral plate of gonarcus large, reniform; 
arch broad. Acumen of tignum long, rounded 
(not swollen) apically. 

Genitalia? (Figs 56-58). Callus cercus ovate; 
28 trichobothria. Subgenitale with short basal 
projection. Spermatheca narrow; ventral impres- 
sion deep; duct long. 

Remarks. C. mediterranea is apparently closely 
related to carnea and shares several characters 
with that species. Like carnea the gena is marked 
with a brown stripe, the wing venation is green, 
costal setae short and intramedian cell short. 
Similarly, the male genitalia of both species have 
many characters in common: spinellae are 
absent, the arcessus is dorsally striated and there 
are few gonosetae. However, the wings are nar- 
rower in mediterranea and the basal dilation of 
the claw much smaller than in carnea and other 
related species. On the basis of the these charac- 
ters it seems justified to continue to treat mediter- 
ranea as a distinct species. 

Duelli (1987) described a population of green 
lacewings referrable to mediterranea from Swit- 
zerland. The population occurred sympatrically 
with carnea but was ecologically separated since 
carnea was confined to oak whereas mediterranea 
was found on pine. In the laboratory the taxa 
interbred and produced fertile hybrids. 

Cianchi & Bullini (1992) report that their 
electrophoretic work on this Swiss population of 
mediterranea suggests it is genetically distinct 
from other populations of mediterranea 
(although they do not state which populations 
they examined) and should be considered a dis- 
tinct taxon. However, until comparable data is 
available from a wide range of populations of this 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



155 



species it is premature to formally recognise the 
taxa as distinct species. 

Distribution. North-west Africa, south-west 
Europe, Hungary (Szentkiralyi, pers comm.), 
Switzerland (Duelli, 1987). 

Material examined (2cf, 1?) Algeria, main- 
land Spain, Majorca. 

Chrysoperla nipponensis (Okamoto) stat. 
rev. 

(Figs 59-67) 

Chrysopa nipponensis Okamoto, 1914: 65. 
Lectotype a", JAPAN: Kyushyu, Kumamoto, 
4.iv.l907 (Kawamura) [Lectotype designation by 
Kuwayama, 1966: 135. ](HU) [examined]. 

Chrysopa kurisakiana Okamoto, 1914: 71. 
Lectotype $, JAPAN: Mt Fuji, 23. vi. 1911 [Lec- 
totype designation by Kuwayama, 1966: 135.] 
(HU) [not examined]. [Synonymized by 
Kuwayama, 1962: 1 1.] 

Chrysopa ilota Banks, 1915: 629. Holotype $, 
PHILIPPINE IS: Luzon, Mt Makiling {Baker) 
(MCZ) [examined]. Syn. n. 

Chrysopa kolthoffiNavds, 1927b: 3. Lectotype 
$, CHINA: Kiangsu (Kolthoff) [Lectotype desig- 
nation by Leraut, 1991: 79.] (MNHN) [exam- 
ined]. [Synonymized with carnea by Tjeder, 
1936: 32; raised to valid species by Leraut, 1992: 
63.] Syn. n. 

Chrysopa kolthoff i var. adaptata Navas, 1934a: 
2. Holotype $, CHINA: Jiangsu Prov., Hufu 
(Loubou), 3.viii.l933 (IZAS) [not examined]. 
Syn. n. 

Chrysopa sinica Tjeder, 1936: 29. Holotype 
Cf, CHINA: South Kansu, Ardjuan, 2900m, 
8.ix.l930 (Sjostedt) (NRS) [examined]. Syn. n. 

Chrysopa (Chrysoperla) sinica Tjeder; Tjeder, 
1966: 391. 

Anisochrysa (Chrysoperla) sinica (Tjeder); 
Holzel, 1970: 51. 

Chrysoperla nipponensis (Okamoto); Tsuk- 
aguchi, 1985: 504 [as synonym of Chrysoperla 
carnea (Stephens)]. 

Anisochrysa (Chrysoperla) sinica (Tjeder); 
Holzel, 1970: 51. 

Chrysopa (Chrysoperla) sinica Tjeder; Tjeder, 
1966: 391. 

Fore wing (Fig. 59) 0\ 10.5-13 mm; $>, 
12.5 mm. Head marked with black/brown stripe 
on gena and lateral clypeus; lateral frons some- 
times marked with red stripe. Maxillary palps 
black dorsally. Antennae shorter than fore wing. 
Pronotum marked with median yellow longitudi- 
nal band and pale brown lateral stripe; protho- 



racic setae long or short, pale. Claw (Figs 60, 61) 
with large basal dilation, about half length of 
claw hook (2.07-2.42: 1; mean 2.22). Fore wing 
relatively broad (length: breadth = 2.9-3.1: 1); 
apex pointed; costal setae long (1.0-2.4 mm), 
inclined towards wing apex; cell im short, not 
meeting 1st Rs crossvein; 1st Rs crossvein usually 
at right-angles to Rs; 4-6 inner gradates, 6-7 
outer gradates; gradates black; crossveins dark at 
each end except costal and Psm crossveins which 
are unmarked at the wing margins. 

Genitaliao" (Figs 62-64). Dorsal apodeme of 
tergite 9 long, extending beyond callus cercus; 
callus cercus broad, ovate; 27-33 trichobothria. 
Membranous apex of sternite 8 + 9 broad, 
rounded. Spinellae absent. Gonosaccus with 
minute spicules; gonosetae few, 5-7 in lateral 
clump. Arcessus with dorsal striations; apex 
blunt, curved ventrally at 90°. Entoprocessus 
large, crescentic. Gonarcus lateral plate broadly 
reniform; arch broad. Acumen of tignum long, 
narrow, rounded apical ly. 

Genitalia9 (Figs 65-67). Callus cercus round; 
32 trichobothria. Subgenitale with short, narrow 
basal extension. Spermatheca with shallow ven- 
tral impression; duct long; vela long. 

Remarks. C. nipponensis is closely related to 
carnea and the male genitalia of the two species 
appear to be indistinguishable. However, there 
are consistent differences in the external mor- 
phology that suggest they should be treated as 
distinct species. In nipponensis the gradates are 
black, the costal setae relatively long and the 
basal dilation of the claw is about half the length 
of the claw hook. In addition, the gena and 
lateral clypeus is heavily marked dark brown/ 
black, the prothoracic setae are pale and the 
acumen of the tignum is relatively long (Fig. 63). 
In carnea the gradates are green, costal setae 
generally short and the basal dilation of the claw 
is about one-third the length of the claw hook. In 
most examples of carnea examined the genal 
markings are less extensive and paler, the pro- 
thoracic setae are black and the acumen is 
shorter. 

Yang & Yang (1990a) examined the male type 
of Chrysopa chusanina Navas and commented 
that the genitalia indicated the species should be 
transferred to Chrysoperla. They also considered 
that the species may be a synonym of C. sinica 
Tjeder. Unfortunately, I have not been able to 
borrow the type of C. chusanina so cannot con- 
firm this view. However, the original description 
of C. chusanina (Navas, 1933c) does not 
resemble that of sinica Tjeder. Navas describes 



156 



S.J. BROOKS 



the palps as yellow, a red arcuate stripe in front 
of the antennae, a red lateral stripe on the head 
and a red lateral stripe on the pronotum. None of 
these markings occur in sinica. However, without 
access to the type material it is impossible to 
decide the true affinities of the species. 

Thierry et al., (1992) and Leraut (1992) incor- 
rectly refer to a French morph of carnea as 
kolthoffi. 

Geographical variation. Korean specimens 
and some of the Chinese material examined have 
a red stripe on the lateral frons that is absent in 
Japanese material. 

The Mongolian paratypes of sinica have the 
gena marked with a narrow, pale brown stripe 
and in one specimen the gradate crossveins were 
green, not marked black. 

Distribution. China, Japan, Korea, Mongolia, 
Philippines. 

Material examined (9d\ 15$) China, Japan, 
Korea, Mongolia. 

Chrysoperla renoni (Lacroix) 

(Figs 68-75) 

Chrysopa renoni Lacroix, 1933: 149. Lecto- 
typecf, FRANCE: Deux-Sevres, Amure, 
29.vi.1930 {Lacroix) [Leraut, 1991: 72. Lecto- 
type designation.] (MNHN) [examined]. 

Chrysoperla ankylopteryformis Monserrat & 
Diaz-Aranda,1989: 261. Holotype 0", SPAIN: 
Almeria, Rambla de Morales (Cabo de Gata), 
10m, 12.viii.1988 [MONSERRAT COLL] 
[examined]. Syn. n. 

Fore wing (Fig. 68) d\ 10.5 mm; $, 11.5 mm. 
Head marked with brown stripe on gena and 
lateral clypeus. Maxillary palps unmarked or 
black dorsally. Antenna very short, little more 
than half length of fore wing. Pronotum marked 
with median longitudinal white/yellow band, 
vague broad pale reddish medio-lateral red 
stripe, narrow antero-ventro-lateral black stripe; 
prothoracic setae very long, pale; setal follicles 
sometimes marked black. Claws with minute 
basal dilation (10.25: 1), claw tooth elongate 
(Fig. 69). Fore wing variable in width (length: 
breadth = 2.8-3.7: 1); rounded apically; vena- 
tion mainly green with costal crossveins some- 
times black at subcosta; costal setae long 
(1.8-2.4 mm), erect; cell im short, not reaching 
1st Rs crossvein; 1st Rs crossvein meets Rs 
obliquely; 4-6 inner gradates, 5-6 outer gra- 
dates. 

Genitaliao" (Figs 70-72). Dorsal apodeme of 



tergite 9 long, extending well beyond callus cer- 
cus; callus cercus broad, ovate; 30 trichobothria. 
Lip of sternite 8+9 narrow, prominent. Spinellae 
absent. Gonosetae few, 5 arranged in lateral 
clump. Arcessus broad with dorsal striations; 
apex blunt, curved at 90°. Entoprocessus large, 
crescentic. Gonarcus with broad lateral plate; 
arch broad. Acumen of tignum broad, not swol- 
len apically. 

Genitalia? (Figs 73-75). Callus cercus broad, 
ovate; 26 trichobothria. Subgenitale with short, 
broad basal extension. Spermatheca with shallow 
ventral impression; duct long; vela short. 

Remarks. On the basis of the male genitalia, C. 
renoni belongs to the carnea-group of species. I 
could detect no useful characters in the gonarcus- 
complex to distinguish renoni from other species 
in this group. However, the acumen of the 
tignum is shorter and broader in renoni than in 
males of carnea. In females of renoni the basal 
lobe of the subgenitale is longer than usual in 
other members of the carnea-group. From other 
Chrysoperla species renoni can be readily identi- 
fied by the long prothoracic and costal setae and 
the minute basal dilation of the claw. The claws 
of C. mediterranea also have a minute basal 
dilation but the claw hook is much shorter than in 
renoni and not at right angles to the claw base. In 
addition, the fore wings of mediterranea are 
much narrower than in renoni and are pointed 
apically rather than rounded. 

The present synonymy between C. renoni and 
C. ankylopteryformis confirms the suggestions of 
Leraut (1991) and Aspock (1992). 

Distribution. France, Spain. 

Material examined (3d\ 3$, 1 without abdo- 
men) 

France (including lectotype of renoni), Spain 
(holotype and two paratypes of ankylopteryfor- 
mis). 

Chrysoperla oblita (Holzel) 

(Figs 76-83) 

Anisochrysa (Chrysoperla) oblita Holzel, 
1973:342. Holotype C\ NEPAL: Jiri, 
12. vi. 1964 (Remane) (ZSBZ) [examined]. 

Chrysoperla oblita (Holzel); Brooks & Barnard, 
1990: 271. 

Fore wing (Fig. 76) cf, 10.5-11 mm; $, 
10.5-13 mm. Head marked with broad reddish 
band across gena, lateral clypeus and labrum, 
markings sometimes extending onto lateral frons 
and medio-lateral area of clypeus; red post- 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



157 



ocular spot. Maxillary palps unmarked. Anten- 
nae as long as fore wing. Pronotum with median 
longitudinal yellow band and red spot in anterior 
corner; prothoracic setae long or short, pale. 
Claw (Fig. 77) with large basal dilation, about 
half length of claw hook (1.88-2.14: 1; mean 
2.01). Fore wing with gradates black; other cross- 
veins dark each end, except Psm crossveins 
entirely green; upper branch of forking marginal 
veins black; anal veins green; apex of wing 
rounded; costal setae long (1.0-1.2 mm), erector 
slightly inclined towards wing apex; 5-7 inner 
gradates, 6-8 outer gradates; cell im long, 1st Rs 
crossvein meets im at apex or subapically; 1st Rs 
crossvein leaves Rs at right-angles or obliquely. 

Genitalias (Figs 78-80). Lip of sternite 8+9 
broad, truncate, at 45° to sternite. Apodeme of 
tergite 9 long, extending beyond callus cercus; 
callus cercus broad, ovate; 28-30 trichobothria. 
Spinellae absent. Gonosaccus with minute spi- 
cules; gonosetae few, 9-12 in lateral clump. 
Arcessus with dorsal striations; apex blunt, 
curved evenly ventrad. Entoprocessus large, 
crescentic. Gonarcus broad, lateral plate foot- 
shaped; arch broad. Acumen of tignum short, 
very broad, rounded apically. 

Genitalia? (Figs 81-83). Callus cercus 
rounded; 25 trichobothria. Subgenitale concave 
basally with median projection. Spermatheca 
with shallow ventral impression; duct long; vela 
short. 

Remarks. C. oblita belongs to the group of 
species related to carnea that have a striated 
arcessus and lack spinellae in the male genitalia. 
However, the species differs from others in the 
group by having relatively more gonosetae in the 
male genitalia {carnea has only 5-6 in each lateral 
group), the gena marked red instead of brown 
and a large basal dilation of the claw. C. oblita is 
readily identifiable by the black gradate cross- 
veins, long costal setae and the broad acumen in 
the male genitalia. 

The holotype of oblita is now in poor condi- 
tion. Both fore wings are broken off at the base. 
The apex of the abdomen is preserved in a vial of 
glycerine but the internal genitalic components 
(gonarcus, arcessus and tignum) are all missing. 
However, Holzel's (1973) figures and desciption 
of the type are sufficient for positive identifica- 
tion of the species. 

Distribution. Highlands of northern Thailand, 
Nepal and north-west India. All specimens were 
collected between 1380 and 1700m. 

Material examined (4q\ 8$) . Thailand: 



Chiang Mai Province. India: Assam, Khasia 
Hills. Nepal. 

Chrysoperla annae sp. n. 

(Figs 84-89) 

Fore wing (Fig. 84) q\ 13.5 mm; 9 unknown. 
Head marked with reddish brown stripe on gena 
and lateral clypeus, red stripe on frontal suture. 
Maxillary palp black dorsally. Antenna shorter 
than fore wing. Pronotum with yellow longitudi- 
nal median band; setae short, black. Claw (Fig. 
85) with small basal dilation, one-quarter or less 
than claw hook length (3.25-9.75: 1; mean 6.5). 
Fore wing narrow (length: breadth = 3.1-3.3: 1), 
venation green; costal setae moderately long 
(0.8-1.0 mm), inclined towards wing apex; 1st Rs 
crossvein meets im subapically; 6-7 inner gra- 
dates, 7-9 outer gradates. 

Genitalia^ (Figs 86-89). Dorsal apodeme of 
tergite 9 short; callus cercus broadly ovate; 25-28 
trichobothria. Lip of sternite 8+9 broad, trun- 
cate when viewed ventrally. Spinellae absent. 
Gonosaccus with 12-20 gonosetae in lateral 
group. Arcessus narrow, straight, striated dor- 
sally; apex rounded, straight or curved ventrad at 
90°. Entoprocessus small, narrow, crescentic. 
Lateral plates of gonarcus narrow, reniform; 
arch broad. Acumen of tignum moderately long, 
narrow not swollen apically. 

Remarks Chrysoperla annae belongs to the 
carnea-group of species. However, the new spe- 
cies possesses several distinctive characters that 
distinguish it from other species in the group. In 
the male genitalia the arcessus, entoprocessus 
and the lateral plates of the gonarcus are nar- 
rower than in carnea and there are 12-20 gonose- 
tae, instead of six, in each lateral group. The 
relatively narrow fore wing with rounded apex is 
also characteristic of the species. 

Distribution. Burma, Tibet. 

Material examined (2cf ) Holotype d\ Tibet: 
Tsaidam (Prejevdoky) (BMNH). Paratype d\ 
Burma: Tongkruk, 9,000, 20.vii.1931 (Kingdon 
Ward) (BMNH). 

Chrysoperla gallagheri Holzel 

(Figs 90-98) 

Chrysoperla gallagheri Holzel, 1989a: 60. Holo- 
type d\ OMAN: Jabal Shams, 2500m, 23.15N/ 
57.15E, 16.vii.1987 {Gallagher) (HOLZEL 
COLL.) [examined]. 



158 



S.J. BROOKS 



Fore wing (Fig. 90) C\ 9.5-10.5 mm; ?, 12 mm. 
Head marked with broad red stripe across gena, 
lateral frons, lateral clypeus; red post-ocular 
stripe; head sometimes suffused entirely red; 
lateral scape and pedicel with red stripe. Maxil- 
lary palp unmarked. Antenna about same length 
as fore wing. Pronotum unmarked, pale median 
longitudinal stripe absent but yellow/green 
median stripe present on pteronotum; protho- 
racic setae long, black; follicles unmarked. Claw 
(Fig. 91) with small basal dilation, less than a 
quarter length of claw hook (5.5: 1). Abdomen 
marked with dorsal median yellow stripe on 
tergites 1-3. Fore wing (length: breadth = 
2.9-3.0: 1), pointed apically; costal setae rela- 
tively long (1.0 mm), inclined towards wing 
apex; 4—5 inner gradates, 5-7 outer gradates; 
gradates black; crossveins black each end; 1st Rs 
crossvein meets cell im at apex; 1st Rs crossvein 
straight or oblique. 

GenitaliacT (Figs 92-95). Lip of sternite 8+9 
narrow, rounded apically; apex of sternite 8+9 
narrowed to form prominent tubercle. Apodeme 
of tergite 9 long, extending far beyond callus 
cercus; callus cercus moderately narrow, ovate; 
26 trichobothria. Spinellae absent. Gonosetae 
numerous, formed into lateral group. Arcessus 
with dorsal striations; apical membrane long, 
narrow, curved at 45°, rounded at tip. Entopro- 
cessus short, narrow, crescentic. Acumen of 
tignum short, broad, swollen; arms of tignum 
narrow. 

Genitalia9 (Figs 96-98). Callus cercus ovate; 
33 trichobothria. Subgenitale with short, narrow 
basal extension. Spermatheca with deep ventral 
impression; duct long; vela short. 

Remarks. Like C. pudica and mutata, the gra- 
dates of gallagheri are black. However, gallagh- 
eri may be distinguished from these species by 
the small basal dilation of the claw, the red stripe 
on the scape and, in males, by the absence of 
spinellae. 

Distribution. Oman. 

Material examined (2d\ 2$) Oman. 

Chrysoperla comans (Tjeder) 
(Figs 99-108) 

Chrysopa (Chrysoperla) comans Tjeder, 1966: 
408. Holotype Q\ ZIMBABWE: Umtali, 
14. i. 1945 (Wright)(TM) [not examined]. 

Chrysoperla comans (Tjeder); Holzel, 1989b: 
175. 

Chryoperla manselli Holzel, 1989b: 176. Holo- 



type cf , SOUTH AFRICA: Transvaal, Vrede- 
huis gardens of Union Buildings, 25.45S/ 
28.12E, l.x.1979, at light (Mansell) (NCI) 
[examined]. Syn. n. 

Fore wing (Fig. 99) cf, 10.5-11.5 mm; $, 
12.0-13.5 mm. Head usually marked with red 
lateral stripe on gena and clypeus; red post- 
ocular stripe. Maxillary palps unmarked. Anten- 
nae about as long as fore wing. Pronotum 
marked with red spot in anterior corner and 
yellow median, longitudinal band; prothoracic 
setae long (2.2-3.0 mm), pale. Claw (Fig. 100) 
with large basal dilation, about two-thirds length 
of claw hook (1.59-1.71: 1; mean 1.65). Fore 
wing broad, with brown gradates; costal setae 
long (2.2-3.0 mm), erect; 5-8 inner gradates, 6-9 
outer gradates; 1st Rs crossvein meets im apically 
or sub-apically. 

Genitaliao" (Figs 101-105). Sternite 8+9 elon- 
gate; lip small, broad. Callus cercus small, nar- 
row, ovate; 26 trichobothria. Apodeme of tergite 
long, often branched, extending considerably 
beyond callus cercus. Spinellae absent. Gonose- 
tae short, numerous, evenly dispersed across 
gonosaccus. Arcessus narrow, apical membrane 
long, blunt, slightly curved or recurved ventrad, 
not striated dorsally. Entoprocessus small cres- 
centic. Lateral plate of gonarcus ovate or boot- 
shaped. Acumen of tignum short, grossly 
swollen. 

Genitalia9 (Figs 106-108). Callus cercus 
ovate; 26 trichobothria. Subgenitale with short 
basal projection. Spermatheca with deep ventral 
impression; duct long; vela long. 

Remarks. C. comans belongs to the group of 
Afrotropical broad-winged species with long, 
erect costal setae. From nyerina and plicata it can 
be distinguished by the absence of spinellae. C. 
comans can be distinguished from the Madagas- 
can species, decaryana, by the narrow arcessus, 
short, swollen acumen and the narrow, elongate 
apex of sternite 8+9. In decaryana the arcessus is 
broad, the acumen narrow and sternite 8+9 is 
not elongated apically. 

When he described manselli, Holzel (1989b) 
remarked on the similarities between manselli 
and comans, in particular the elongation of stern- 
ite 8+9, which does not occur elsewhere in the 
genus. However, he then went on to say that the 
gonarcus complex and tignum were totally differ- 
ent in the two taxa. Nevertheless, on 
re-examination of the types of manselli I can see 
no significant differences in the genitalia from 
that described by Tjeder (1966) for comans. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



159 



Indeed Holzel's (1989b) figure of the acumen of 
manselli shows it less swollen than it is in fact. 

Distribution. Kenya, Zimbabwe, South 
Africa. 

Material examined (3cf, 39) ■ South Africa, 
Kenya. 



Chrysoperla decaryana (Navas) 
(Figs 109-114) 

Chrysopa decaryana Navas, 1934b: 60. Holo- 
typecf, MADAGASCAR: Tananarive, 1921 
(Decary) (MNHN) [examined]. 

Chrysoperla decaryana (Navas); Holzel. 1989b: 
169. 

Fore wing (Fig. 109) d\ 13 mm; 9 ur| known. 
Head marked with broad red lateral stripe on 
gena, frons and clypeus; red post-ocular stripe. 
Maxillary palp with black dorsal stripe. Antenna 
shorter than fore wing. Pronotum marked with 
yellow/white median longitudinal band and red 
medio-lateral spot; prothoracic setae long, pale. 
Basal dilation of claw large, about half length of 
claw hook (2.25: 1) (Fig. 110). Fore wing broad 
(length: breadth = 3: 1); costal setae long 
(1.6 mm), slightly inclined apically; Rs crossveins 
sinuate; gradates black and other crossveins dark 
at each end except costals at costa and Psc 
crossveins entirely green; 1st Rs crossvein meets 
im at apex. 

Genitalias (Figs 111-114). Lip of sternite 8+9 
narrow, tapering apically, prominent; callus cer- 
cus narrow, ovate; 32 trichobothria; apodeme of 
tergite 9 short, not extending beyond callus cer- 
cus, branched. Spinellae absent. Gonosaccus 
with minute spicules. Gonosetae few (about 14) 
in median group. Entoprocessus minute, crescen- 
tic. Arcessus lacking dorsal striations; short, 
broad, straight; apex blunt, rounded. Gonarcus 
with broad arch; basal dilation narrow wedge. 
Acumen of tignum long, parallel sided, rounded 
apically; arms narrow. 

Remarks. C. decaryana belongs to the group of 
Afrotropical species with broad wings and long 
costal setae. It is probably most closely related to 
comans Tjeder since both species lack spinellae 
in the male genitalia. However, from this species 
decaryana can be distinguished by the broad 
arcessus and relatively narrow acumen. 



Chrysoperla siamensis sp. n. 

(Figs 115-123) 

Fore wing (Fig. 115) cf, 11-12.5 mm; $, 14 mm. 
Head sometimes unmarked or marked with red- 
dish spot on lateral clypeus, extending to medio- 
lateral area; red spot on lateral frons which 
sometimes extends posteriorly as stripe between 
eye and scape; red post-ocular stripe; gena 
unmarked, or sometmes with red spot. Maxillary 
palps unmarked. Antennae shorter than fore 
wing. Pronotum marked with yellow median 
longitudinal band, bordered yellow/orange, 
sometimes marked with red medio-lateral spot; 
prothoracic setae short, pale. Claw (Fig. 116) 
with large basal dilation, over half length of claw 
hook (1.86: 1). Fore wing broad (length: breadth 
= 2.8-3.0: 1); rounded apically; costal setae long 
(1.0-1.4 mm), erect; 1st Rs crossvein leaves Rs at 
oblique angle; 1st Rs crossvein meets cell im 
subapically; Rs crossveins sinuous; gradates 
green or greyish with surrounding membrane 
suffused grey, other veins green; 4-7 inner gra- 
dates, 6-8 outer gradates. 

Genitalias (Figs 117-120). Lip of sternite 8+9 
broad, rounded. Dorsal apodeme of tergite 9 
extending beyond callus cercus; callus cercus 
broad, ovate; 23-38 trichobothria. Spinellae 
absent. Gonosetae numerous in median group. 
Arcessus lacking dorsal striations; narrow, curv- 
ing in apical half to 90°; membranous tip very 
long and narrow, blunt apically. Entoprocessus 
very small, narrow, crescentic (not visible in 
some specimens examined). Gonarcus with lat- 
eral plates broad, ovate; arch broad. Acumen of 
tignum short, swollen apically. 

Genitalia? (Figs 121-123). Callus cercus 
ovate; 28 trichobothria. Subgenitale with narrow 
basal extension. Spermatheca with shallow ven- 
tral impression; duct short; vela short. 

Remarks. C. siamensis is unusual amongst 
Chrysoperla species in lacking both spinellae and 
dorsal striations on the arcessus in the male 
genitalia. The only other species that also lack 
these characters are comans, decaryana and gal- 
lagheri. Other distinctive characters of siamensis 
are the broad, rounded fore wing with a grey 
suffusion around the gradate crossveins, the rela- 
tively long intramedian cell, sinuous Rs cross- 
veins and short pale prothoracic setae. In 
addition, the arcessus is very narrow and the lip 
of sternite 8+9 unusually broad. 

Distribution. Thailand. 



Material examined (1 cf). Madagascar. 



Material examined (5$, 2$) Holotype cf, 



160 



S.J. BROOKS 



Thailand: Chiang Mai Prov., Mae Rim to 
Samoeng Road, 24 Km post, 1100m, 29.U989 
(Cotton) (BMNH). Paratypes. Thailand: 2cf, 
Korat Prov., Khao Yai National Park, Khao 
Kieow, 1200m, l.iii.1989 (Cotton); 1$, Chiang 
Mai Prov., Doi Suthep-Pui National Park, 
1380m, 24.ii. 1989 (Cotton); lcf, 1$, Doi Suthep- 
Pui National Park, 1460m, 26.iv-10.v.l989 (Cot- 
ton); lcf, Doi Suthep-Pui National Park, 1200m, 
10-11. xi. 1988 (Bradley, Lewvanich, Fletcher) 
(BMNH). 

Chrysoperla nyerina (Navas) 
(Figs 124-132) 

Chrysopa nyerina Navas, 1933a: 210. Holotype 
d\ KENYA: Nyeri, 9.vi.l932 (MNHN) 
[examined]. 

Chrysopa socia Navas, 1936a: 116. Lectotype $, 
KENYA: Marakwet, Elgeyo Escarpment, 
2500m, 1932-33 (Arambourg, Chappuis & 
Jeannel) [Lectotype designation by Holzel, 
1989b: 173.] (MNHN) [examined]. Syn. n. 

Chrysoperla nyerina (Navas); Holzel, 1989b: 
172. 

Chrysoperla socia (Navas); Holzel, 1989b: 173. 

Fore wing (Fig. 124) cf, 13-14.5 mm; $, 
13.5-15 mm. Head occasionally unmarked or 
marked with broad red lateral stripe on gena and 
clypeus; red post-ocular spot. Maxillary palps 
black dorsally, sometimes unmarked. Specimens 
examined from Uganda had a red lateral stripe 
on the scape, otherwise scape unmarked. Prono- 
tum with longitudinal yellow median band, 
sometimes with red spot in anterior corner; pro- 
thoracic setae very long, pale. Claw (Fig. 125) 
with large basal dilation, about two-thirds length 
of claw hook (1.77-1.89: 1; mean 1.83). Fore 
wing broad (length: breadth = 2.6-2.8: 1); costal 
setae long (1.8-2.6 mm), erect; gradates black 
and other crossveins dark at each end (except Psc 
crossveins which are green at wing margin); 6 
inner gradates, 6-7 outer gradates; im long, 1st 
Rs crossvein meets im sub-apically. Abdomen 
with pair of red dorsal spots on basal segments. 

Genitalias (Figs 126-129). Lip of sternite 8+9 
broadly rounded. Apodeme of tergite 9 long, 
extends far beyond callus cercus; callus cercus 
broad, ovate; 21-36 trichobothria. Few (about 
20) minute spinellae, each spine on an individual 
tubercle. Gonosetae numerous, evenly distrib- 
uted on gonosaccus. Entoprocessus small, cres- 
centic. Arcessus long, narrow with dorsal 
striations; weakly curved ventrad; truncate api- 
cally. Gonarcus with narrow lateral plate. Acu- 



men of tignum long, parallel sided, rounded 
apically. 

Genitalia? (Figs 130-132). Callus cercus 
ovate; 25 trichobothria. Subgenitale broad. Sper- 
matheca narrow; ventral impression deep; vela 
short; duct long. 

Remarks. C nyerina is probably closely related 
to plicata since this is the only other species in the 
genus with simple spinellae. However, there are 
several characters that will distinguish the spe- 
cies. In nyerina the basal dilation of the claw is 
larger than in plicata. In the male genitalia the 
apex of the acumen is not swollen in nyerina and 
it has fewer and smaller spinellae than plicata. In 
plicata, unlike nyerina, the spinellae are not set 
in tubercles. 

Distribution. Kenya, Sudan, Uganda. 

Material examined (4cf , 6$) Kenya, Sudan, 
Uganda 

Chrysoperla plicata (Tjeder) 
(Figs 133-141) 

Chrysopa (Chrysoperla) plicata Tjeder, 1966: 
406. Holotype cf, ZIMBABWE: Marandellas, 
Eirene Farm, x.1960 (NHMZ) [examined]. 

Chrysopa plicata (Tjeder); Holzel, 1989b: 168. 

Fore wing (Fig. 133) cf, 9.5-12.5 mm; $, 
12.5-13 mm. Head marked with strong red spot; 
red post-ocular stripe. Maxillary palps 
unmarked. Antennae about as long as fore wing. 
Pronotum marked with red stripe in each corner 
and red centro-lateral spot; median longitudinal 
yellow band; prothoracic setae pale, long or 
short. Claw (Fig. 134) with large basal dilation, 
about two-thirds length of claw tooth (1.58: 1); 
rounded posteriorly. Fore wing marked with 
black spot on tympanal organ; crossveins black at 
each end (except marginal crossveins at wing 
margin); gradates black; 3-7 inner gradates, 4-8 
outer gradates; costal setae long (1.0—1.8 mm), 
slightly inclined; im variable in length. 

Genitalias (Figs 135-138). Apodeme of terg- 
ite 9 variable, usually not extending beyond 
callus cercus; 21-30 trichobothria; lip of sternite 
8 + 9 prominent, strongly tapered apically when 
viewed ventrally. Spinellae present, numerous, 
single broad spines not mounted on tubercle. 
Gonosetae numerous, evenly distributed across 
gonosaccus. Arcessus with dorsal striations; 
blunt apically, weakly curved. Entoprocessus 
short, narrow, triangular. Gonarcus with wide 
arch and median projection; lateral plates broad, 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



161 



reniform. Acumen of tignum short, broad, bul- 
bous apically. 

Genitalia? (Figs 139-141). Callus cercus 
rounded; 29 trichobothria. Subgenitale with 
large basal extension. Spermatheca narrow; ven- 
tral impression shallow; vela short; duct long. 

Remarks. Males of C. plicata are unusual 
amongst Chrysoperla species in that the arcessus 
is striated and spinellae are present. The only 
other species to share these characters is nyerina. 
C. plicata also shares with nyerina simple spinel- 
lae in the male genitalia. Unlike nyerina, how- 
ever, the spinellae of plicata arise directly from 
the gonosaccus and are not raised on tubercles. 
In most other species in the genus which possess 
spinellae, these structures are composed of sev- 
eral spines positioned on each tubercle. C. pli- 
cata may be distinguished from other 
Chrysoperla species by the strongly defined red 
spot on the gena and the three red lateral stripes 
on the pronotum. In addition, the claw is 
strongly reflexed and has a large basal dilation. 

Tjeder's (1966) figure of the male genitalia of 
plicata shows the apex of the arcessus strongly 
hooked. In the specimens examined during this 
study all had the arcessus only weakly curved. 
Unfortunately, the genitalia of the holotype have 
been slide mounted in such a way that it is 
impossible to see the extent of the curvature of 
the arcessus in the holotype. 

Distribution Afrotropics. 

Material examined (9o", 3$) Mozambique, 
Sene-Gambia, Somalia, South Africa, Zaire, 
Zambia, Zimbabwe. 

Chrysoperla insulata (Fraser) 
(Figs 142-148) 

Chrysopa insulata Fraser, 1957: 26. Syntypesd" 
$, REUNION: Hellbourg; forest track in the 
forest of Belouve; Bebour & Plaine des Caf- 
res, Piton Manuel, I. 1955 (Paulian) (IRSM & 
BMNH) [ld\ 1$ syntype in BMNH exam- 
ined]. 

Chrysoperla insulata (Fraser); Brooks & Bar- 
nard, 1990: 271. 

Fore wing (Fig. 142) o\ 13 mm; $, 16.5 mm. 
Head marked with black lateral stripe on 
clypeus. Maxillary palps black. Antennae shorter 
than fore wing. Pronotum with median longitudi- 
nal yellow band; prothoracic setae short, pale. 
Claw (Fig. 143) with large basal dilation, over 
half the length of claw hook (2.21: 1). Fore wing 
broad (length: breadth = 28.-2.9: 1), pointed 



apically; costal setae long (2.4-2.8 mm), erect; 
im long, apex extends beyond 1st Rs crossvein; 
gradates black; 8-9 inner gradates, 9-10 outer 
gradates; other crossveins darkened at each end. 

Genitalias (Figs 144-145). Lip of sternite 8+9 
broad, rounded; apodeme of tergite 9 short, not 
extending beyond callus cercus; 31 trichobothria. 
Spinellae present, numerous, 2-3 spines on 
tubercle. Gonosetae numerous, arranged in lat- 
eral group. Arcessus narrow; lacking dorsal stria- 
tions; strongly curved ventrad; apical membrane 
long, narrow, truncate at apex. Entoprocessus 
minute, narrow, straight. Gonarcus narrowly 
expanded basally. Tignum absent. 

Genitalia? (Figs 146-148). Callus cercus 
ovate; 39 trichobothria. Subgenitale broad with 
narrow basal extension. Spermatheca with mod- 
erate ventral impression; vela short; duct long. 

Remarks. C. insulata is unique amongst 
Chrysoperla species because the tignum is absent 
in the male genitalia. The species is also charac- 
terized by the relative lack of head markings, 
only the clypeus is marked with a lateral black 
stripe, and the broad distinctly pointed fore 
wings. C. insulata is the largest Chrysoperla 
species with a fore wing length of 16.5 mm in 
females. 

Distribution. Reunion. 

Material examined (1q\ 1 9 syntypes) 
Reunion. 

Chrysoperla pudica (Navas) 
(Figs 149-159) 

/Hetnerobius brevicollis Rambur, 1842: 427. 
Holotype sex unknown, Mauritius (Marchal) 
[not traced]. [Nomen dubium] 

Chrysopa pudica Navas, 1914a: 82. Holotype O", 
SOUTH AFRICA: Transvaal, Piet Retief 
(Crawshay) (BMNH) [examined]. 

Chrysopa incongrua Navas, 1914c: 97. Holotype, 
sex unknown, SENEGAL-NIGER: Koulikoro 
(NAVAS COLL.) [type not traced]. Provi- 
sional synonym. 

Chrysopa nigriciana Navas, 1931b: 119. Holotype 
$, NIGERIA: Guezzan & Air, Prince Sixte de 
Bourbon, 1929 (MNHN) [examined]. Syn. n. 

Chrysopa (Chrysoperla) pudica (Navas); Tjeder, 
1966: 397. 

Chrysoperla pudica (Navas); Ohm & Holzel, 
1982: 160. 

Chrysoperla nigricana (Navas); Brooks & Bar- 
nard, 1990: 271. 



162 



S.J. BROOKS 



Fore wing (Fig. 149) d\ 8.5-12 mm; 9, 
11-13 mm. Head markings variable, unmarked 
or sometimes marked with red post-ocular spot; 
red spot on gena; some specimens with short, 
black lateral stripe on clypeus. Maxillary palps 
unmarked. Antennae as long as fore wing. 
Pronotum with median yellow longitudinal 
stripe; often with red spot in each corner or 
medio-lateral spot; prothoracic setae pale, long 
or short. Claw (Fig. 150) with large basal dila- 
tion, about two-thirds length of claw hook 
(1.57-1.76: 1; mean 1.7). Fore wing with black 
gradates; crossveins in basal half of wing black at 
each end; 3-6 inner gradates, 4-9 outer gradates; 
im short, not meeting 1st Rs crossvein or apex of 
im meets crossvein; costal setae short (0.6 mm), 
inclined towards wing apex. 

Genitalias (Figs 152-156). Apodeme of terg- 
ite 9 short, not extending beyond callus cercus; 
callus narrow, ovate; 26-35 trichobothria; lip of 
sternite 8+9 broad rounded; sternite 8+9 with 
subapical ventral swelling. Spinellae numerous, 
6-8 spines arranged on tubercle. Gonosetae 
numerous, evenly distributed on gonosaccus. 
Arcessus lacking dorsal striations; apical mem- 
brane short, rounded, abruptly curved, some- 
times reflexed. Entoprocessus short, narrow. 
Lateral plates of gonarcus broadly reniform; 
gonarcus arch broad. Acumen of tignum long, 
narrow, swollen apically. 

Genitalia? (Figs 157-159). Callus cercus 
ovate; 37 trichobothria. Subgenitale heart- 
shaped with broad basal extension. Spermatheca 
with deep ventral impression; vela short; duct 
long. 

Remarks. Holzel & Ohm (1991; 1992) identi- 
fied a small Chrysoperla species from Mauritius 
as Chrysoperla brevicollis (Rambur). I have 
examined this material and it is the same as C. 
pudica. Unfortunately, the type of brevicollis is 
missing so the identity of this species must be 
gleaned from the original description. Rambur's 
(1842) description is brief and in my opinion it is 
impossible to say with certainty the genus, let 
alone the species, referred to in this description. 
One of the most characteristic features of the 
Mauritian Chrysoperla species is its black gradate 
crossveins. However, in Rambur's description 
the venation is said to be very pale yellowish 
green and no mention is made of dark veins. In 
view of this I think that it is preferrable to treat 
Hemerobius brevicollis as nomen dubium rather 
than as a senior synonym of Chrysoperla pudica. 
Holzel & Ohm (1992) treat the populations of 
pudica in Mauritius and mainland Africa as dis- 



tinct species. However, I was unable to find any 
consistent character to separate the two taxa. In 
general specimens of pudica from Mauritius (fore 
wing length 9.5-10.5 mm) are smaller than those 
from the mainland. However, the smallest speci- 
men that I examined during this study was a male 
from Nigeria with a fore wing length of 8.5 mm. 

Specimens of pudica can be distinguished by 
the pale maxillary palps, large basal dilation of 
the claw and dark gradates. Males possess tuber- 
culate spinellae and lack dorsal striations on the 
arcessus. C. pudica has not been recorded from 
Madagascar (Holzel & Ohm, 1992) but this may 
simply be due to under collecting rather than 
disjunct distribution. C. mutata McL. is close to 
pudica and the species have many characters in 
common. However, the basal dilation of pudica 
is considerably larger than mutata. In mutata the 
basal tooth is less than half the length of the claw 
hook but in pudica the basal tooth is two-thirds 
the length of the claw hook. Also the angle 
between the basal tooth and the claw hook is 
greater in mutata than pudica. 

I was unable to trace the type of Chrysopa 
incongrua Navas but it seems likely that the 
species is synonymous with pudica. Navas 
(1914c) describes the species as close to congrua 
with a black stripe on the lateral clypeus and 
yellow palps, typical markings for West African 
specimens of pudica. Nevertheless, he does not 
mention any darkening of the gradate crossveins 
which casts doubt on this interpretation. 

Geographic variation. West African speci- 
mens are marked with a short black fleck later- 
ally on the clypeus. Southern African specimens 
are marked with a red post-ocular spot between 
the eye and vertex. 

Distribution. Throughout Afrotropics. 

Material examined (17 d\ 16 $) Ascension 
I., Mozambique, Namibia, Nigeria, South 
Africa, St Helena, Senegal, Zambia, Zimbabwe. 

Chrysoperla mutata (McLachlan) 
(Figs 160-170) 

Chrysopa mutata McLachlan, 1898: 167. Lecto- 
type cf, ALGERIA: Biskra, 30.iii.1895 
(Eaton) [Lectotype designated by Kimmins, 
1940: 447] (BMNH) [examined]. 

Chrysopa nepia Navas, 1911: 266. Holotype Q, 
ALGERIA (MNHN) [examined]. Syn. n. 

Chrysopa libera Navas, 1928: 463. LECTOTYPE 
Cf [here designated], LIBYA: Cyrenaica, 
Oasis Giarabub, 16.iii.1927 (Canfalonieri) 
(MCSN) [examined]. Syn. n. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



163 



Chrysopa quettana Navas, 1931a: 83. Lectotype 
^.PAKISTAN: Baluchistan, Quetta, 

21. v. 1930 (MNHN) [Lectotype designated by 
Leraut, 1991: 79.] [examined]. [Synonymized 
with carnea by Aspock et «/., 1980: 268.] Syn. 
n. 

Chrysopa phaeocephala Navas, 1931b: 120. 
Holotype 0\ TUNISIA: Bou Medma 
(Dumont) (MNHN) [examined]. Syn. n. 

Chrysopa grazianii Navas, 1932: 418. LECTO- 
TYPE Cf [here designated], LIBYA: Cufra, 
v.1931 (MCSN). [examined]. Syn. n. 

Chrysopa expurgata Tjeder, 1949: 83. Holotype 
Cf, ISRAEL: Kinneret, -200m, lO.v.1936 (Pal- 
moni) (PALMONI COLL., ISRAEL) [not 
examined]. [Synonymized by Aspock et at., 
1980: 270.] 

Chrysopa (Chrysoperla) expurgata Tjeder; 
Tjeder, 1966: 391. 

Chrysopa (Chrysoperla) tnutata McLachlan; H61- 
zel, 1967: 35. 

Anisochrysa (Chrysoperla) mutata (McLachlan); 
Holzel, 1970:51. 

Anisochrysa (Chrysoperla) concinna Holzel, 
1974: 44. Holotype d\ Iran: Bandarabbas, 
3-10.iv.1972 (Ressl) (ASPOCK COLL.) 
[examined]. Syn. n. 

Chrysoperla mutata (McLachlan); Canard & 
Laudeho, 1978: 71. 

Chrysoperla concinna (Holzel); Holzel, 1980: 
169. 

Chrysoperla nepia (Navas); Brooks & Barnard, 
1990:271. 

Chrysoperla phaeocephala (Navas); Brooks & 
Barnard, 1990: 271. 

Fore wing (Fig. 160) cf, 9-10.5 mm; $, 
9.5-12.5 mm. Head marked or unmarked on 
gena with faint red spot; short narrow black 
stripe on lateral clypeus. Maxillary palp 
unmarked. Antennae as long as fore wing. 
Pronotum marked with longitudinal median yel- 
low band; sometimes with dorso-lateral median 
red/brown spot or greyish antero-lateral spot; 
prothoracic setae long or short, pale. Basal dila- 
tion of claw about half length of claw hook 
(1.71-2.64: 1; mean 2.29) (Fig. 161-163). Fore 
wing crossveins dark at each end, except mar- 
ginal veins at wing margin unmarked; gradates 
black; tympanal organ often marked with black 
spot; costal setae short (0.6 mm), inclined 
towards wing apex; 1-5 inner gradates; 2-6 outer 
gradates; cell im variable in length. 

Genitalias (Figs 164-167). Lip of sternite 8+9 
small, rounded; sternite 8+9 with small subapical 
swelling. Apodeme of tergite 9 short, not extend- 



ing beyond callus cercus; callus cercus narrow, 
ovate; 23-33 trichobothria. Spinellae numerous, 
each tubercle bearing 5-7 spines. Gonosetae 
numerous, about 25 in lateral group. Arcessus 
lacking dorsal striations; apical membrane long, 
bent ventrally at 45-90°, tip pointed. Entoproces- 
sus narrow, crescentic. Gonarcus with lateral 
plate broad, reniform; arch broad. Acumen of 
tignum long, narrow or swollen apically. 

Genitalias (Figs 168-170). Callus cercus 
rounded; 33 trichobothria. Subgenitale heart- 
shaped. Spermatheca with moderate ventral 
impression; vela short; duct long. 

Remarks C. mutata is most similar to puclica 
but can be distinguished by the shape of the claw. 
In mutata the basal dilation of the claw is about 
half the claw hook length, the posterior margin is 
relatively straight and the angle between the 
basal tooth and the claw hook is relatively large 
whereas in puclica the basal dilation is more than 
half the claw hook length, the posterior margin is 
curved and the angle between the basal tooth and 
the claw hook is less than in mutata. 

Distribution From north Africa through Ara- 
bia and the Middle East to Pakistan and northern 
India. 

Material examined (26cf, 24$) Algeria, 
Egypt, India (Punjab, Rajastan), Iran, Iraq, 
Kuwait, Mauritania, Saudi Arabia, Spain, Tuni- 
sia. 

Chrysoperla nigrinervis sp. n. 

(Figs 171-178) 

Fore wing (Fig. 171) d\ 9 mm; 9< 10.5 mm. 
Head marked with red spot on gena; long, broad 
black stripe on lateral clypeus. Maxillary palps 
unmarked. Antennae slightly longer than fore 
wing. Pronotum marked with longitudinal 
median yellow band; greyish lateral stripe; pro- 
thoracic setae short, pale. Claw (Fig. 178) with 
very small basal dilation, less than one-quarter 
length of claw hook (5.8: 1). Fore wing costal 
setae short (0.6 mm), inclined towards wing 
apex; crossveins entirely black; longitudinal 
veins black adjacent to crossveins; tympanal 
organ marked with black spot; gradates black; 
3-4 inner gradates, 5-6 outer gradates; cell im 
short not reaching 1st Rs crossvein or meets at 
apex. Abdominal tergites marked with large 
black spot on posterior margin. 

Genitalias (Figs 172-174). Lip of sternite 8+9 
small, rounded; sternite 8+9 with small subapical 
swelling. Apodeme of tergite 9 short, not extend- 



164 



S.J. BROOKS 



ing beyond callus cercus; callus cercus narrow, 
ovate; 31-32 trichobothria. Spinellae numerous, 
each tubercle bearing 5-7 spines. Gonosetae 
numerous, about 20 in lateral group. Arcessus 
lacking dorsal striations; apical membrane long, 
bent ventrally at 45°, tip pointed. Entoprocess 
small, narrow, crescentic. Gonarcus with broad 
lateral reniform plate; arch broad. Acumen of 
tignum long, narrow, slightly swollen at apex. 

Genitalia9 (Figs 175-177). Callus cercus 
ovate; 37 trichobothria. Subgenitale with long 
narrow basal extension. Spermatheca with mod- 
erate ventral impression; duct long; vela long. 

Remarks. The male genitalia of nigrinervis are 
apparently indistinguishable from mutata but 
there are sufficient external characters to regard 
specimens from the Canary Islands as a distinct 
species. C. nigrinervis can be distinguished from 
mutata and the closely related pudica by the 
more extensively blackened venation, head, pro- 
thoracic and abdominal markings together with 
the small basal dilation of the claw. In addition, 
in females of nigrinervis the subgenitale has a 
narrower basal lobe than specimens of mutata 
and pudica which have a heart-shaped subgeni- 
tale. 

Distribution. Canary Islands. 

Material examined (2d\ 3$) Holotype d\ 
Canary Islands: Gran Canada, Maspalomas, 
14.viii.1966 (Guichard & Ward) (BMNH) 
Paratypes lo", same data as holotype; 3$, same 
data as holotype, 17-28. vi. 1966 (BMNH) 

Chrysoperla exul (McLachlan) 
(Figs 179-187) 

Chrysopa exul McLachlan, 1869: 23. Holotype 
0\ ST HELENA (Melliss) (BMNH) [exam- 
ined]. 

Chrysopa wollastoni Navas, 1914a: 80. Holotype 
(apex of abdomen missing), ST HELENA, 
1877 (Wollaston) (BMNH) [examined]. [Syn- 
onymized by Kimmins, 1940: 446]. 

Chrysoperla exul (McLachlan); Holzel, 1989b: 
176. 

Fore wing (Fig. 179) d\ 12-13 mm, $, 
13-14.5 mm. Head marked with red spot on 
outer angle of frons; red lateral spot on clypeus; 
red spot on lateral vertex; red spot between base 
of antennae; scape with faint lateral red stripe; 
maxillary palps black dorsally. Antennae longer 
than fore wing. Pronotum marked with brown 
spot in each corner and red cento-median spot; 
median yellow longitudinal band; prothoracic 



setae short, pale. Claw (Fig. 180) with large basal 
dilation, about two-thirds length of claw hook 
(1.74: 1). Fore wing quite broad (length: breadth 
= 2.7-3.0: 1); costal setae short (0.4-0.6 mm); 
venation entirely green; 6-8 inner gradates, 7-9 
outer gradates; im variable in length. 

Genitalias (Figs 181-184). Lip of sternite 8+9 
short, broad; callus cercus broad ovate; 33 tri- 
chobothria; apodeme of tergite 9 branched, 
extending only a short way beyond callus cercus. 
Spinellae present, 6-7 spines on numerous small 
tubercles. Gonosetae numerous. Entoprocessus 
small, crescentic. Arcessus straight, extreme 
apex curved at 45°; lacking dorsal striations; 
apical membrane of arcessus long, narrow, trun- 
cate apically. Gonarcus with broad reniform lat- 
eral plate; arch narrow. Acumen of tignum long, 
narrow. 

Genitalia9 (Figs 185-187). Callus cercus 
ovate; 38 trichobothria. Subgenitale with narrow 
basal extension. Spermatheca with shallow ven- 
tral impression; vela short; duct long. 

Remarks. C. exul may be distinguished from 
other species of Chrysoperla by the characteristic 
head and pronotal markings, in particular the 
brown spot in each corner of the pronotum and 
the red spots on the vertex and between the 
antennae. 

The possession of synapomorphies in exul and 
congrua suggests a close relationship between 
these two species. In both these species the 
gonarcus arch is narrow and the spinellae are 
tuberculous. 

Distribution. St Helena. 

Material examined (6d\ 4$) St Helena. 

Chrysoperla congrua (Walker) 
(Figs 188-195) 

Chrysopa congrua Walker, 1853: 238. Holotype 
(abdomen missing), WEST AFRICA 
(BMNH) [examined]. 

Chrysopa concolor Walker, 1853: 239. Holotype 
(abdomen missing), ZAIRE (BMNH) [exam- 
ined]. [Synonymized by Tjeder, 1966: 392.] 

Chrysopa otalatis Banks, 1910a: 102. Holotype 
$, AUSTRALIA: Queensland, Brisbane, 
vi.1904 (MCZ) [not available for examina- 
tion]. Syn. n. 

Chrysopa bequaerti Navas, 1912: 409. Lectotype 
d\ ZAIRE: Kongolo, 9.ii.l911 (Bequaert) 
[lectotype designation by Tjeder, 1966: 393.] 
(MRAC) [examined]. [Synonymized by 
Esben-Petersen, 1928a: 218.] 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



165 



Chrvsopa lemoulti Lacroix, 1923: 119. Holotype 
Cf, NEW CALEDONIA (MNHN) [exam- 
ined]. [Synonymized with otalatis by Esben- 
Petersen, 1928b: 103.] Syn. n. 

Chrvsopa clypealis Navas, 1929b: 103. Holotype 
$, ZAIRE: Haut-Uele, Yebo Moto, Abmiva. 
1925 {Burgeon) (MRAC) [examined]. Syn. n. 

Chrysopa solaria Navas, 1930: 411. Holotype 0\ 
EAST AFRICA: Marangu (ZMH) 
[destroyed]. [Synonymized by Holzel, 1989b: 
166.] 

Chrysopa bequaerti var. occipitalis Navas, 1931c: 
127. Holotype $, ZAIRE (MRAC) [exam- 
ined]. Syn. n. 

Chrysopa congrua var. rubella Navas, 1931c: 
127. Holotype $, ZAIRE (MRAC) [exam- 
ined]. Syn. n. 

Chrvsopa acutella Navas, 1933a: 209. Holotype 
Cf .MADAGASCAR: Majunga (MNHN) 
[examined]. [Synonymized by Holzel, 1989b: 
166.] 

Chrysopa (Chrysoperla) congrua (Walker); 
Tjeder, 1966:392. 

Chrysoperla congrua (Walker); Semeria & Qui- 
lici, 1986: 110. 

Chrysoperla otalatis (Banks); Brooks & Barnard, 
1990:271. 

Chrysoperla acutella (Navas); Brooks & Bar- 
nard, 1990: 271. 

Chrvsoperla clypealis (Navas); Brooks & Bar- 
nard, 1990:271. 

Fore wing (Fig. 188) O" 10-12 mm; 9 
12-13.5 mm. Head usually marked with red lat- 
eral stripe on gena, clypeus, sometimes extend- 
ing to frons; red post-ocular spot often present. 
Maxillary palp unmarked or marked black dor- 
sally. Antenna about as long as fore wing. Prono- 
tum with longitudinal median yellow band and 
often with red or greyish spot in anterior corner; 
prothoracic setae long, pale (short in Easter I. 
specimens). Claw (Fig. 189) with small basal 
dilation, less than one quarter length of claw 
hook (4.1-6.25: 1; mean 5.05). Fore wing narrow 
(length: breadth = 3.0-4.2: 1), pointed apically; 
5-8 inner gradates, 5-9 outer gradates; im very 
short; costal setae moderately long 
(0.8-1.0 mm), inclined; veins green except cl 
and c2 crossveins maybe darkened; 1st Rs cross- 
vein at right-angles to Rs. 

Genitalias (Figs 190-192). Lip of sternite 8+9 
short, narrow, downcurved. Callus cercus broad, 
ovate; 24-33 trichobothria; apodeme of tergite 9 
extends beyond callus cercus. Spinellae sparse, 
4-6 spines arranged on small tubercles. Gonose- 
tae numerous, evenly distributed across gonosac- 



cus. Arcessus narrow, weakly curved, lacking 
dorsal striations, apex blunt and slightly swollen. 
Entoprocessus very small, short and narrow. 
Gonarcus with narrow arch and narrow, wedge- 
shaped lateral plates. Acumen of tignum short, 
bulbous. 

Genitalia9 (Figs 193-195). Callus cercus 
rounded. Basal extension of subgenitale usually 
long and narrow. Spermatheca small with, deep 
ventral impression; vela short; duct long. 

Remarks. C. congrua is characterized by pos- 
sessing green gradates, a small basal dilation of 
the claw and narrow fore wings with a short 
intramedian cell. A close relationship between 
congrua and the Neotropical species of Chrysop- 
erla is suggested by the synapomorphous posses- 
sion of a swollen apex to the arcessus and 
bulbous acumen (cf. C. externa). 

From New's (1980) description of Chrysopa 
itatotis Banks, evidently based on the female 
holotype. it is apparent that the species is refer- 
able to Plesiochrysa (comb. n.). However, the 
figure he gives of the male genitalia are those of a 
Chrysoperla species. On examination of the eight 
specimens New (1980) identified as C. italotis, 
from the ANIC, it is apparent that in fact these 
are misidentfied specimens of Chrysoperla con- 
grua. 

Geographical variation. Most material 
examined from Australia, New Caledonia and 
New Hebrides had the maxillary palps 
unmarked. In specimens from Samoa, Easter 
Island and West Malaysia the palps were always 
black dorsally. The palps were variable in speci- 
mens from other regions. All the Sri Lankan 
specimens were collected between 4,000 and 
7,000 feet. 

Distribution. Throughout Australasia from 
the Pacific Islands to Sri Lanka (absent from 
India), throughout the Afrotropics and recorded 
from Oman on the Arabian Peninsular (Holzel & 
Ohm, 1992) 

Material examined (117o\ 196$). Afrotro- 
pics: Angola, Botswana, Ethiopia, Fernando Po, 
Gabon, Ghana, Ivory Coast, Kenya, Madagas- 
car, Malawi, Mozambique, Nigeria, Prinicipi 
Island, Ruanda, South Africa, Tanzania, 
Uganda, Urundi, Zaire, Zambia, Zimbabwe. 
Oriental: Sri Lanka, Thailand, West Malaysia. 
Australasian: Australia, New Caledonia, New 
Hebrides, Papua New Guinea. Pacific: Easter I., 
Fiji, Rapa I., Samoa. 



166 



S.J. BROOKS 



Chrysoperla dozieri (Smith) 
(Figs 196-202) 

Chrysopa dozieri Smith, 1931: 808. Holotype G\ 
HAITI: Port au Prince, 29. ii. 1929 (Smith) 
(MCZ) [examined]. 

Chrysoperla dozieri (Smith); Brooks & Barnard, 
1990: 271. 

Fore wing (Fig. 196) 0\ 10.5-12 mm; §, not 
seen. Head marked with red L-shaped stripe 
across frontal suture; red stripe on gena and 
lateral clypeus; narrow medio-lateral red spot on 
front of vertex. Maxillary palps marked black 
dorsally. Antennae as long as fore wing. Prono- 
tum marked with longitudinal median yellow 
band; medio-lateral black spot and usually with 
red stripe in anterior corner; prothoracic setae 
short, pale. Claw (Fig. 197) with large basal 
dilation, about half length of claw hook (2.08: 1). 
Fore wing pointed at apex; costal setae short 
(0.4—0.6 mm), inclined towards wing apex; 5-6 
inner gradates, 6-8 outer gradates; gradates 
black; other crossveins black each end; cell im 
long, 1st Rs crossvein meets im at apex or 
sub-apically; 1st Rs crossvein straight or oblique. 

GenitaliacT (Figs 198-202). Lip of sternite 8+9 
broad, rounded. Apodeme of tergite 9 forked 
above and below callus, short; callus cercus 
broad, ovate; 29 trichobothria. Spinellae few, 
tuberculate. Gonosetae numerous, in lateral 
group. Arcessus broad, straight; dorsal striations 
absent; membranous tip spade-like (narrows sub- 
apically, then abruptly widens at apex), truncate 
at apex, curved ventrad at 90°. Entoprocessus 
not visible. Gonarcus lateral plates broadly trian- 
gular; arch broad. Acumen of tignum narrow, 
hardly swollen apically. 

Remarks. This species can be readily distin- 
guished from other species in the genus by the 
presence of a pair of narrow red spots on the 
vertex. Some Cuban specimens of exotera may 
also have a pair of red spots on the front of the 
vertex but they are broader than those of dozieri. 
Unlike dozieri, specimens of exotera do not have 
a red stripe below the fronto-clypeal suture. The 
spade-like apex of the arcessus in the male geni- 
talia is also distinctive. C. dozieri differs from 
other Chrysoperla species in the rufilabris-group 
in plesiomorphically retaining a broad gonarcus 
arch and a broad arcessus. The species also has 
fewer spinellae tubercles than other rufilabris- 
group species. C. dozieri is endemic to Haiti and 
this geographical isolation may account for its 
apparent phylogenetic isolation. 



Distribution. Haiti. 

Material examined (2d\ 2 examples with 
abdomens missing) Haiti. 

Chrysoperla defreitasi sp. n. 

(Figs 203-208) 

Fore wing cf (Fig. 203), 11 mm; $ unknown. 
Head marked with broad red stripe on gena, 
lateral clypeus and lateral labrum; narrow red 
stripe on lateral fronto-clypeal suture. Maxillary 
palps marked black dorsally. Pronotum with yel- 
low longitudinal median stripe and narrow 
brownish red lateral stripe; prothoracic setae 
long, pale. Claw (Fig. 208) with large basal 
dilation, about two-thirds length of claw hook 
(1.68: 1). Fore wing relatively narrow (length: 
breadth = 3: 1); venation mostly green with 
black gradates, basal costal crossveins entirely 
black, black spot at each end of radial and Rs 
crossveins, c,/c 2 crossvein entirely black; mar- 
ginal branches of veins Cu 2 , 1A and 2 A entirely 
black; apex of fore wing pointed; costal setae 
short (0.6 mm), inclined towards wing apex; im 
short, not reaching 1st Rs crossvein; 5 inner 
gradates, 7 outer gradates. 

GenitaliacT (Figs 204-207). Dorsal apodeme of 
tergite 9 long; callus cercus narrow ovate; 28-29 
trichobothria; lip of sternite 8 + 9 short, broad. 
Spinellae present, 29 large tubercles with 7-8 
spicules. Gonosetae numerous 50+ in lateral 
group. Arcessus broad, tapering subapically 
before widening to form spatulate apex; dorsal 
striations absent. Entoprocessus narrow, cres- 
centic. Gonarcus with broad, wedge-shaped lat- 
eral plates; arch relatively broad, truncate 
medially with bilobed median projection. Acu- 
men of tignum long, narrow, slightly swollen at 
apex. 

Remarks. C. defreitasi closely resembles C. 
externa and C. exotera. From C. exotera, C. 
defreitasi cannot be reliably distinguished on 
external characters although the claw hook is 
larger in defreitasi than in exotera. However, the 
male genitalia are quite different. In C. defreitasi 
the arcessus is broad with a spatulate apex 
whereas in C. exotera the arcessus is narrow and 
truncate at the apex. 

C. defreitasi may be distinguished from externa 
by the black gradate crossveins, which are green 
in externa, and the basal dilation of the claw, 
which is considerably larger in defreitasi than 
externa. Certain apomorphic charcters in the 
male genitalia suggest that C. defreitasi may be 
closely related to C. dozieri, from Haiti. In both 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



167 



species the unstriated arcessus is broader than is 
typical in neotropical species of the genus, the 
apex of the arcessus is spatulate and the gonarcus 
arch is relatively broad. Although the entopro- 
cessus are narrow in C. defreitasi, they are larger 
than other Chrysoperla species in species-group 
4.2 (see p. 143). 

Distribution. Brazil. 

Material examined (Id") Holotype cf, Bra- 
zil: Jaquaritingas, 15. xi. 1992 (de Freitas) (USP). 

Chrysoperla asoralis (Banks) 
(Figs 209-217) 

Chrysopa asoralis Banks, 1915: 627. Holotype 
0\ PERU: Chosica, 2800', 7.vi. (Parish) 
(MCZ) [examined]. 

Chrysopa oscillans Navas, 1922a: 260. LECTO- 
TYPE Cohere designated], PERU: Arequipa, 
1912 (Escomel) (MNHN) [examined]. Syn. n. 

Chrysopa lindana Navas, 1924b: 111. Holotype 
$, PERU: on the Chilean frontier, 1921 (Mar- 
tin) (MNHN) [examined]. Syn. n. 

Chrysopa arequipae Navas, 1929a: 18. Holotype 
$,PERU: Arequipa, 1922 (Escomel) 
(MNHN) [examined]. Syn. n. 

Chrysoperla asoralis (Banks); Brooks & Bar- 
nard, 1990:271. 

Fore wing (Fig. 209) d\ 10-14 mm, $. 
12-16 mm. Head marked with red stripe on 
gena, lateral clypeus and frons, occasionally with 
vertex red dorsally; red post-ocular spot. Maxil- 
lary palps black dorsally. Antennae as long as 
fore wing. Pronotum with median longitudinal 
yellow stripe; red spot in anterior corner; protho- 
racic setae short, pale. Claw (Fig. 210) with large 
basal dilation, over half length of claw hook 
(1.83-1.87: 1; mean 1.85). Fore wing pointed at 
apex; costal setae short (0.6-0.8 mm), inclined 
towards wing apex; cell im variable in length; 1st 
Rs crossvein meets Rs at right-angles; 5-8 inner 
gradates, 6-9 outer gradates; gradates black; 
other crossveins dark at each end, except Psm 
crossveins unmarked. 

Genitalias (Figs 214-217). Lip of sternite 8 + 9 
broad, rounded. Dorsal apodeme of tergite 9 
long, extending well-beyond callus cercus; callus 
cercus narrow, ovate; 20-41 trichobothria. 
Spinellae numerous, each tubercle with 4-5 long 
spines. Gonosetae numerous, 50+ in lateral 
group. Arcessus narrow, arcuate, occasionally 
recurved apically; dorsal striations absent; apical 
membrane long, pointed at tip. Entoprocessus 
minute, slender, crescentic. Gonarcus arch nar- 



row; narrow wedge-like lateral plate. Acumen of 
tignum long, swollen at apex. 

Genitalia9 (Figs 211-213). Callus cercus 
rounded; 32 trichobothria. Subgenitale with nar- 
row basal extension. Spermatheca small with 
moderate ventral impression; vela moderate; 
duct long. 

Remarks. C asoralis most closely resembles 
exotera and ruflabris, both of which have black 
gradate crossveins and black maxillary palps. 
From exotera, asoralis can be distinguished by 
the red post-ocular spot, which is absent in 
exotera, and the apical membrane of the arcessus 
which is pointed at the tip in males of asoralis but 
truncate and slightly swollen in males of exotera. 
C. rufilabris also differs from asoralis in the 
shape of the apex of the arcessus which is much 
longer and is abruptly narrowed in the former 
species. In addition, the acumen is shorter and 
broader in rufilabris than asoralis. The spinellae 
are unusually long in asoralis. 

The type of lindana differs from other speci- 
mens of asoralis in having a longer fore wing 
(16 mm, which was 2 mm longer than the next 
largest specimen) and the vertex marked red with 
a narrow median interruption. However, I con- 
sider these differences to be due to individual 
variation and not of taxonomic significance. 

Distribution. Neotropics. 

Material examined (20cf, 149) Argentina, 
Bermuda. Guatemala, Peru, Venezuela. 

Chrysoperla exotera (Navas) 
(Figs 218-226) 

Chrysopa exotera Navas, 1914a: 87. Holotype $, 
GUATEMALA: Duenas, Capetillo (Cham- 
pion) (BMNH) [examined]. 

Chrysopa extranea Navas, 1922b: 170. Holotype 
Cf .CUBA (MCZ) [examined]. Syn n. 

Chrysopa estradai Navas, 1924a: 334. Holotype, 
CUBA: Habana, Rio Almendares, 
23-26.X.1923 (Cervera) (HAVANA) [not 
examined]. Syn. n. 

Chrysopa torrei Navas, 1924a: 335. Holotype cf , 
CUBA: Habana, Rio Almendares, 8.ix.l923 
(Cervera) (MCZ) [examined]. Syn. n. 

Chrysopa barbouri Navas, 1924a: 335. Holotype 
sex unknown (abdomen missing), CUBA: 
Habana, Rio Almendares, 8.xii.l923 (Cervera) 
(MCZ) [examined]. Syn. n. 

Chrysopa tenera Navas, 1924a: 336. Holotype $, 
CUBA: Habana, Rio Almendares, 8.xii.l923 
(MCZ) [examined]. Syn. n. 



168 



S.J. BROOKS 



Chrysopa exterior Navas, 1925: 190. [Replace- 
ment namefor C. extranea Navas]. 

Chrysopa cubensis Navas, 1927a: 75. [Unneces- 
sary replacement name for C. extranea Navas]. 

Chrysopa incisa Banks, 1948: 163. Holotype £, 
MEXICO: Chihuahua, Ciudad Juarez, 
4.vi.l927 (Dampf) (MCZ) [not examined, not 
traced in MCZ]. Syn. n. 

Chrysopa quadornia Banks, 1948: 164. Holotype 
$, MEXICO: Acapulco, 19.xii.1924 {Dampf) 
(MCZ) [not examined, not traced in MCZ]. 
Syn. n. 

Chrysopa castalia Banks, 1948: 165. Holotype $, 
MEXICO: Tamaulipas, Santa Engracia, 
3.ii.l936 {Dampf) (MCZ) [not examined, not 
traced in MCZ]. Syn. n. 

Chrysoperla exotera (Navas); Brooks & Barnard, 
1990: 271. 

Fore wing (Fig. 218) cf, 13 mm; 9, 
11.5-13.5 mm. Head marked with red spot on 
gena, lateral clypeus and lateral frons; anterior 
margin of vertex sometimes with red stripe or 
paired spots; entire head occasionally suffused 
red. Maxillary palps black dorsally. Antennae 
shorter than fore wing. Yellow longitudinal band 
on pronotum present or absent; occasionally with 
red spot in anterior corner; prothoracic setae 
long, pale. Claw (Fig. 219) with large basal 
dilation, about half claw hook length (2.07: 1). 
Fore wing apex pointed; costal setae short 
(0.6-0.8 mm), inclined towards wing apex; gra- 
dates black or greyish; crossveins black at each 
end except at wing margins; 5-7 inner gradates, 
7-9 outer gradates; cell im short, apex not reach- 
ing 1st Rs crossvein; 1st Rs crossvein straight or 
oblique. 

Genitalias (Figs 220-223). Lip of sternite 8+9 
broad, rounded. Dorsal apodeme of tergite 9 
short, forked above or below callus cercus; callus 
cercus broad, ovate; 40 trichobothria. Spinellae 
numerous, about 8 spines on each tubercle. 
Gonosetae numerous, arranged in lateral group. 
Arcessus narrow, arcuate; dorsal striations 
absent; apical membrane short, tip broadened, 
truncate. Entoprocessus minute, narrow, cres- 
centic. Gonarcus with lateral plates narrow, reni- 
form or wedge-shaped; arch narrow. Acumen of 
tignum short, broad, swollen at apex. 

Genitalia 9 (Figs 224-226). Callus cercus 
rounded; 32 trichobothria. Subgenitale with nar- 
row basal extension. Spermatheca narrow with 
deep ventral impression; vela long; duct moder- 
ate. 

Remarks. This species is very similar to 



comanche but can be distinguished by the black 
gradate crossveins which are green in comanche. 
In addition, the apex of the arcessus and acumen 
in exotera is swollen, whereas in comanche they 
are narrow. 

Distribution. Central America and Cuba. 

Material examined (lcf, 4$) Cuba, Costa 
Rica, Guatemala, Mexico. 

Chrysoperla externa (Hagen) 
(Figs 227-236) 

Chrysopa externa Hagen, 1861: 221. Syntypes, 
USA: Washington, California; Mexico (MCZ) 
[not traced]. 

Chrysopa lanata Banks, 1910b: 154. Holotype 
CT, ARGENTINA: Mendoza {Jensen-Haarup) 
(MCZ) [examined]. [Synonymized by Adams, 
1962: 222.] 

Chrysopa graciana Navas, 1919: 301. Holotype, 
ARGENTINA: Alta Gracia, ii.1918 {Bruch) 
(LP) [not examined]. [Synonymized with 
argentina Navas by Adams, 1975: 169. Synony- 
mized with externa by Adams & Penny, 1987: 
421.] 

Chrysoperla externa (Hagen); Adams, 1983: 43. 

Chrysoperla externa cocosensis Adams, 1983: 42. 
Holotype Q\ COSTA RICA: Cocos I., Wafer 
Bay, 9. hi. 1980 {Sherry & Werner) (LACM) 
[examined]. Syn. n. 

Fore wing (Fig. 227) Q\ 10.5-11.5 mm; 9, 
12.5 mm. Head marked with broad red stripe on 
gena and lateral clypeus; red post-ocular stripe. 
Maxillary palps black dorsally. Antennae about 
same length as fore wing. Pronotum with yellow 
longitudinal median band; usually with reddish 
spot in anterior corner; prothoracic setae long or 
short, pale or black. Claw (Fig. 228-229) with 
small basal dilation, less than half claw hook 
length (2.4-2.54: 1; mean 2.48). Fore wing nar- 
row (length: breadth = 3.1-3.5: 1), apex 
pointed; costal setae short (0.6-1.0 mm), 
inclined towards wing apex; 5-7 inner gradates, 
6-8 outer gradates; gradates green; crossveins in 
basal half of wing often darkened at each end; 
cell im very short, not reaching 1st Rs crossvein; 
1st Rs crossvein meets Rs at right angles. 

Genitalias (Figs 230-233). Lip of sternite 8+9 
rounded. Dorsal apodeme of tergite 9 long, 
forked below callus; callus cercus narrow, ovate; 
28-32 trichobothria. Spinellae numerous, each 
tubercle with 5-6 spines. Gonosetae numerous, 
positioned centrally on gonosaccus. Arcessus 
without dorsal striations; membranous apex of 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



169 



arcessus short, bulbous, curved ventrally at 45°. 
Entoprocessus small triangular, often not visible. 
Gonarcus with lateral plates narrow, wedge- 
shaped; arch narrow. Acumen of tignum long, 
slightly swollen apically. 

Genitalia9 (Figs 234-236). Callus cercus 
ovate; 32 trichobothria. Subgenitale with broad, 
rounded basal extension. Spermatheca narrow; 
ventral impression deep; vela short; duct long. 

Remarks. C. externa is similar in appearance to 
comanche but the basal dilation of the claw is 
smaller in externa and the apex of the arcessus is 
narrow in comanche but swollen in externa. C. 
externa appears to be closely related to congrua. 
since the male genitalia are very similar and the 
apex of the arcessus is swollen in both species. 
However, although the apex of the acumen is 
swollen in both externa and congrua, the neck is 
longer in externa. In addition the basal dilation of 
the claw is noticeably larger in externa than 
congrua. 

I have examined a female specimen deposited 
in the MCZ which has been identified as C. 
externa Hagen by Banks (USA: Florida, Winter 
Park, 31.x., Davis). This specimen bears a purple 
label, type written 'neotype'. I have not been 
able to trace any reference to this specimen being 
nominated a neotype so this designation has no 
validity. The specimen has been correctly identi- 
fied but since there is no confusion regarding the 
identity of C. externa and considering that the 
specimen was not collected in the same region as 
the original type series I see no merit in formally 
recognising this specimen as a neotype. 

Adams (1983) distinguished C. e. cocosensis 
from C. e. externa on the basis of the arcessus 
being sclerotised medially in the former taxon 
but sclerotised laterally with a median membra- 
nous area in the latter. However, examination of 
the holotype of C. e. cocosensis revealed that 
Adams had misinterpretted the structure of the 
arcessus and that, in fact, the lateral sclerotised 
strips are separated by a membranous median 
area. The arcessus of other males examined from 
Cocos Island were also indistinguishable from 
typical examples of C. externa. 

Geographical variation. In Mexican speci- 
mens the post-ocular spot is absent, the lateral 
frons marked with red stripe, the basal dilation of 
the claw larger and the intramedian cell longer 
than typical specimens. 

Distribution. Throughout Neotropics, extend- 
ing northwards to Florida and South Carolina 
(Adams, 1962). 



Material examined (27d\ 48$) Argentina, 
Belize, Brazil, Costa Rica (Cocos Island), Guate- 
mala, Mexico, Peru, Uruguay, USA (Florida), 
Venezuela, West Indies (Bahamas, Barbados, 
Cayman, Trinidad). 

Chrysoperla galapagoensis (Banks) 

(Figs 237-245) 

Chrysopa galapagoensis Banks, 1924: 179. Holo- 
type, ECUADOR: Galapagos Is, South Sey- 
mour, 23. iv. 1923 (Beebe) (MCZ) [not 
examined]. 

Chrysoperla galapagoensis (Banks); Brooks & 
Barnard, 1990:271. 

Fore wing (Fig. 237) cf, 9-10.5 mm; $, 
10.5 mm. Head marked with black stripe on gena 
and lateral clypeus; red stripe on lateral frons, 
lateral labrum and dorso-lateral clypeus; red 
post-ocular stripe present or absent. Maxillary 
palps black dorsally. Antennae shorter than fore 
wing. Pronotum usually marked with narrow 
black median stripe and latero-median black 
spot, yellow longitudinal stripe absent; protho- 
racic setae long, black or pale. Claw (Fig. 238) 
with small basal dilation, less than half length of 
claw hook (2.73: 1). Fore wing pointed apically; 
costal setae short (0.6-0.8), inclined towards 
wing apex; 4-6 inner gradates, 5-6 outer gra- 
dates; gradates black; crossveins entirely black, 
except Psc crossveins in apical half of wing; cell 
im short, not quite meeting 1st Rs crossvein or 
crossvein meets apex of cell; 1st Rs crossvein 
oblique or at right-angles to Rs. 

Genitalias (Figs 239-241). Lip of sternite 8+9 
narrow, rounded. Apodeme of tergite 9 short, 
branched below callus cercus; callus cercus nar- 
row, ovate; 23-26 trichobothria. Spinellae few, 
6-12 small tubercles bearing about 4 spines. 
Gonosetae numerous. Arcessus narrow, gently 
curved; lacking dorsal striations; apex rounded 
and slightly expanded. Entoprocessus not visible. 
Gonarcus with lateral plates narrow, wedge- 
shaped; arch narrow. Acumen of tignum short, 
slightly swollen apically. 

GENITALIA9 (Figs 242-245). Callus cercus 
broadly ovate; 28 trichobothria. Subgenitale with 
short, narrow basal extension. Spermatheca with 
shallow ventral impression; vela short; duct 
short. 

Remarks. C. galapagoensis is distinctive 
amongst Chrysoperla species because of the 
extensive black venation, prominent black lateral 
stripe on the head and narrow black median 



170 



S.J. BROOKS 



stripe on the pronotum. The male genitalia are 
also distinctive since the spinellae are few and 
small. The arcessus is slightly swollen apically 
which is reminiscent of externa but the tip of the 
arcessus is more bulbous in externa than galapa- 
goensis. 

Distribution. Galapagos Islands. 

Material examined (5cf, 5$) Galapagos 
Islands (Fernandina, Hood, Indefatigable, 
Jervis, Santiago, South Seymour, Tower). 

Chrysoperla comanche (Banks) 
(Figs 246-255) 

Chrysopa comanche Banks, 1938: 119. Holotype 
$, USA: Texas, Laredo (MCZ) [examined]. 

Chrysopa sperryae Banks, 1943: 74. Holotype $, 
USA: California, Riverside (Sperry) (MCZ) 
[not examined]. [Synonymized by Adams, 
1956: 45.] 

Chrysoperla comanche (Banks); Garland, 1985b: 
738. 

Fore wing (Fig. 246) cf, 8-11 mm; $, 
12-12.5 mm. Head marked with red stripe on 
gena and lateral clypeus, usually extending to 
lateral frons; red post-ocular spot present or 
absent; occasionally head totally unmarked. 
Maxillary palps black dorsally. Antennae about 
as long as fore wing. Pronotum with yellow 
longitudinal median stripe; sometimes with 
dorso-median black spot. Claw (Fig. 247) with 
large basal dilation, over half length of claw hook 
(1.86-2.0: 1; mean 1.93). Fore wing apex 
pointed; costal setae short (0.4—0.6 mm), 
inclined towards wing apex; Rs crossveins in 
stigmatic region sinuate; 3-6 inner gradates, 4—8 
outer gradates; gradates green, sometimes inner 
gradates greyish; crossveins in basal half of wing 
sometimes dark at ends; cell im short, not reach- 
ing 1st Rs crossvein; 1st Rs crossvein oblique or 
at right-angles to Rs. 

Genitalias (Figs 248-252). Lip of sternite 8+9 
broad, rounded. Apodeme of tergite 9 long, 
forked below callus; callus cercus narrow, ovate; 
26-29 trichobothria. Spinellae numerous, 4—5 
long spines on tubercle. Gonosetae numerous, 
50+ in lateral group. Arcessus narrow, arcuate; 
dorsal striations absent; apical membrane short, 
truncate or slightly broadening. Entoprocessus 
minute, slender crescentic. Gonarcus with nar- 
row lateral plates; arch narrow. Tignum with 
acumen long, slightly widened apically. 

Genitalia9 (Figs 253-255). Callus cercus 
rounded; 30 trichobothria. Subgenitale with 



long, narrow basal extension. Spermatheca with 
moderate ventral impression; duct short; vela 
moderate. 

Remarks. C. comanche is very similar to 
exotera but there are two characters that suggest 
they are distinct species. Specimens assignable to 
exotera have black gradate crossveins, swollen 
acumen and the arcessus is markedly swollen at 
the extreme apex. In comanche the gradates are 
green, the acumen is narrow and the apex of the 
arcessus is narrow or slightly widened. The two 
species are also separated geographically, with 
exotera occurring in Central America but 
comanche being restricted to southwest USA. C. 
comanche can be distinguished from most New 
World Chrysoperla species by the green gra- 
dates. From the carnea-group of species 
comanche can be separated by the red stripe on 
the gena since the gena is marked brown in the 
carnea-group. From externa, comanche differs by 
having a large basal dilation of the claw, and 
narrow arcessus. In externa the basal dilation of 
the claw is distinctly less the half the length of the 
claw hook and the arcessus is swollen apically. 

Material examined from the Hawaiian Islands 
had been previously identified as C. lanata Banks 
(Zimmerman, 1957), which is now considered as 
a synonym of C. externa (Hagen) (Adams, 1962). 
However, all Hawaiian specimens examined in 
this study proved to be comanche and there is no 
evidence that externa occurs on the Hawaiian 
Islands. 

Distribution. South-west USA, Hawaii. 

Material examined (51d\ 48$) . USA (Ari- 
zona, California); Hawaiian Islands (Kauai, 
Lanai, Maui, Molokai, Oahu). 

Chrysoperla harrisii (Fitch) 

(Figs 256-265) 

Chrysopa perla Harris, 1841: 197. Holotype, 
USA: New England (MCZ) [not examined]. 
[Homonym of Chrysopa perla L.] 

Chrysopa harrisii Fitch, 1855: 794. [New name 
for C. perla Harris]. 

Chrysopa stenostigma Navas, 1914b: 61. Holo- 
type (sex unknown), USA: New York, Long 
Island, Yaphank, 25.vii.1913 (NAVAS 
COLL.) [not traced]. [Synonymized by Bick- 
ley & MacLeod, 1956: 193.] 

Chrysoperla harrisii (Fitch); Garland, 1985a: 93. 

Fore wing (Fig. 256) cf, 12.5 mm; $, 13-14 mm. 
Head marked with broad red stripe on gena, 
lateral frons and lateral clypeus; broad red post- 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



171 



ocular stripe. Maxillary palps black dorsally. 
Antennae slightly longer than fore wing. Prono- 
tum marked with yellow longitudinal median 
band; brownish spot in anterior corner; protho- 
racic setae short, black or pale; setal follicles 
black. Claw (Fig. 257) with very small basal 
dilation, less than one-quarter claw hook length 
(8: 1). Fore wing narrow (length: breadth = 
3.2-3.4: 1), apex pointed; venation entirely 
green; costal setae short (0.8-1.0 mm), inclined 
towards wing apex; cell im long, 1st Rs crossvein 
meets im at apex or subapically; 1st Rs crossveins 
leaves Rs obliquely; 7-9 inner gradates, 9-10 
outer gradates. 

Genitalias (Figs 258-262). Lip of sternite 8+9 
broad, rounded. Dorsal apodeme of tergite 9 
long, extending far beyond callus cercus; callus 
cercus narrow, ovate; 32 trichobothria. Spinellae 
present, numerous tubercles bearing about 7 
spines. Gonosetae numerous, arranged in lateral 
group. Arcessus without dorsal striations; 
strongly curved ventrad; membranous tip bent at 
90°, truncate apically. Entoprocessus minute, 
crescentic. Lateral plate of gonarcus narrow 
wedge-shaped; arch narrow. Acumen of tignum 
short, broad, slightly swollen apically. 

Genitalia? (Figs 263-265). Callus cercus 
broadly ovate; 37 trichobothria. Subgenitale with 
long, narrow basal extension. Spermatheca with 
shallow ventral impression; vela short; duct 
short. 

Remarks. C. harrisii is one of the narrow 
winged species of Chrysoperla but is character- 
ized by the entirely green venation and extremely 
small basal dilation of the claw. The male genita- 
lia of harrisii are similar to those of comanche 
which suggests that the the two species may be 
closely related. However, these species may be 
differentiated by the head markings, which are 
more extensive in harrisii than comanche and the 
shape of the acumen, which is broader in harrisii 
than comanche. 

Distribution. Canada (Garland, 1982); USA: 
California, Maryland, Michigan, Virginia (Bram 
& Bickley, 1963); Montana, Connecticut; 
Mexico. 

Material examined (2cf, 3$) . USA: Mon- 
tana; Connecticut. 

Chrysoperla mexicana sp. n. 

(Figs 266-274) 

Fore wing (Fig. 266) o", 10.5 mm; $, 11.5 mm. 
Head marked with broad red lateral stripe on 



gena, frons and clypeus with blackish flecks on 
gena and clypeus. Maxillary palp black dorsally. 
Antennae about same length as fore wing; outer 
margin of scape with apical brown stripe; pedicel 
and flagellar segments entirely dark brown. 
Pronotum marked with yellow median longitudi- 
nal band and brownish red lateral stripe; protho- 
racic setae short, black. Claw (Fig. 267) 
undilated basally. Fore wing narrow (length: 
breadth = 3.4-3.6:1), apex pointed; gradate 
crossveins green; costal, radial and Psc cross- 
veins black, darker in basal half of wing; veins 
7/4,2/4, Cu 2 , base of Rs and Psm at cell M 2 and 
im all black; other veins green; costal setae short 
(0.6 mm), inclined towards wing apex; cell im 
short, 1st Rs meets Psm just beyond apex of im; 
1st Rs crossvein leaves Rs at right angles; 5-6 
inner gradates, 7-8 outer gradates. 

Genitalias (Figs 268-271). Lip of sternite 8+9 
broad, rounded. Dorsal apodeme of tergite 9 
short, not extending beyond callus cercus. Callus 
cercus broad, ovate; 29 trichobothria. Spinellae 
present, numerous tubercles with 5-7 spines. 
Gonosetae numerous, arranged in lateral group. 
Arcessus narrow, without dorsal striations; apex 
curved ventrally at 90°; apical membrane moder- 
ately long, truncate at tip. Entoprocessus very 
narrow, crescentic. Lateral plate of gonarcus 
narrow, quadrangular; arch narrow. Acumen of 
tignum narrow, swollen at apex. 

Genitalia? (Figs 272-274). Callus cercus 
broad, ovate; 26 trichobothria. Subgenitale with 
long narrow basal lobe. Spermatheca with shal- 
low ventral impression (everted in figure); vela 
short; duct short. 

REMARKS. Chrysoperla mexicana can be readily 
distinguished by the black antennae, in all other 
species of the genus the antennae are green. 
Other distinguishing characters are the broad 
brownish lateral stripe on the pronotum, the 
undilated base of the claw and the narrow fore 
wings with dark venation in the basal half of the 
wing. 

C. mexicana appears to be closely related to 
harrisii since both species share a number of 
apomorphies including narrow fore wings, claws 
with small basal dilation, truncate apex of arces- 
sus, and long basal lobe on the spermatheca. 

I am most grateful to Prof. P. A. Adams for 
drawing my attention to this species. 

Distribution. Mexico. 

Material examined (ld\ 1$) Holotype $, 
Mexico: Sinaloa, 36.6 miles NE Villa Union, 



172 



S.J. BROOKS 



near La Capilla Detaxte, 2.vii.l982 (SDNHM) 
Paratype 9, same data as holotype. 

Chrysoperla rufilabris (Burmeister) 

(Figs 275-285) 

Chrysopa rufilabris Burmeister, 1839: 979. Syn- 
types, Mid-USA & MEXICO (ZMH) [type 
destroyed]. 

Chrysopa interrupta Schneider, 1851: 76. Holo- 
type d\ USA: Pennsylvania (NMHU) [not 
examined]. [Synonymized by Bram & Bickley, 
1963: 6.] 

Chrysopa attenuata Walker, 1853: 242. Lectotype 
d\ USA: Florida, St John's Bluff (Doubleday) 
[Lectotype designation by Kimmins, 1940: 
447.] (BMNH) [examined]. [Synonymized by 
Banks, 1903: 161.] 

Chrysopa repleta Walker, 1853: 244. Holotype 
(abdomenmissing), USA: Georgia (BMNH) 
[examined]. [Synonymized by Banks, 1903: 
161.] 

Chrysopa novaeboracensis Fitch, 1855: 90. Syn- 
types, USA: New York State (MCZ) [not 
examined]. [Synonymized by Hagen, 1866: 
395.] 

Chrysopa tabida Fitch, 1855: 92. USA (MCZ) 
[not examined]. [Synonymized by Hagen, 
1866: 398]. 

Chrysopa citri Ashmead, 1880: 13. Holotype, 
USA: Florida [not examined]. [Synonymized 
by Bickley & MacLeod, 1956: 194.]. 

Chrysopa medialis Banks, 1903: 154. Holotype 
9, USA: Maryland, Colombia, High Is., ix 
(MCZ) [not examined, not traced in MCZ]. 
[Synonymized by Bram & Bickley, 1963: 6.] 

Chrysoperla rufilabris (Burmeister); Garland, 
1985a: 93. 

Fore wing (Fig. 275) d", 11-12 mm; $, 
11.5-12.5 mm. Head marked with broad red 
stripe on gena, lateral clypeus and lateral frons; 
red post-ocular spot usually present; anterior 
margin of vertex sometimes marked with red 
stripe, which may extend between antennae. 
Maxillary palp black dorsally. Antenna slightly 
longer than fore wing. Pronotum marked with 
yellow longitudinal median stripe; often with red 
spot in anterior corner; prothoracic setae short, 
pale. Claw (Fig. 276-278) with large basal dila- 
tion, over half length of claw hook (1.85-2.23: 1; 
mean 2.02). Fore wing relatively narrow (length: 
breadth = 3.1-3.3: 1), apex pointed; costal setae 
moderately long (0.8-1.2 mm), inclined towards 
wing apex; 5-7 inner gradates, 6-8 outer gra- 
dates; gradates black; crossviens black at each 
end, except costal crossveins at costa and Psc 



crossveins entirely green; cell im long, 1st Rs 
crossvein meets im subapically or at apex; 1st Rs 
crossvein meets Rs obliquely or at right angles. 
Abdomen green with yellow median longitudinal 
stripe on tergites and dorso-lateral red spot on 
tergites 1 and 2; in some specimens (possibly 
showing winter colouration) abdomen dark 
brown dorsally, posterior corner of tergite black- 
ish brown, pale brown ventrally. 

Genitalias (Figs 279-282). Lip of sternite 8+9 
broad, rounded; with subapical patch of microse- 
tae. Dorsal apodeme of tergite 9 short, extending 
a little beyond callus cercus; callus cercus nar- 
row, ovate; 24-32 trichobothria. Spinellae 
present, numerous small tubercles bearing 6-7 
spines. Gonosetae numerous, arranged in lateral 
group. Arcessus narrow; dorsal striations absent; 
apical membrane long, narrow, pointed at tip, 
curved ventrad at 90°; 3^4- mid-lateral microsetae 
present or absent. Entoprocessus minute or not 
visible, narrow crescentic. Gonarcus with lateral 
plates narrow, wedge-shaped; arch narrow. Acu- 
men of tignum short, broad, rounded apically. 

Genitalia9 (Figs 283-285). Callus cercus 
rounded; 27 trichobothria. Subgenitale with 
long, narrow basal extension. Ventral impression 
of spermatheca variable, shallow or deep; vela 
long; duct long. 

Remarks. The black gradate crossveins will 
serve to distinguish C. rufilabris from all other 
North American Chrysoperla species except 
some specimens of C. johnsoni and C. downesi. 
However, spinellae are absent in the male genita- 
lia of C. johnsoni and the gena is often marked 
with a brown stripe. In C. rufilabris spinellae are 
present in the male genitalia and the gena is 
invariably marked red. The long, narrow mem- 
branous apex of the arcessus in males of rufila- 
bris is unique within the genus. 

Geographic variation. Male specimens from 
Bermuda, Florida, Louisiana, South Carolina, 
Texas and Mexico were found to have 3-4 short 
setae situated laterally, mid-way along the arces- 
sus. The setae were absent in males examined 
from Canada, Florida, Georgia, North Carolina, 
Oklahoma, Virginia. Up to 11 such setae were 
also reported by Bram & Bickley (1963) in 82% 
of specimens examined from Louisiana and 15% 
from Maryland, and in specimens from Kansas, 
Texas and Florida. Garland (1982) did not find 
any Canadian specimens with the setae. I was 
unable to find any secondary characters that 
conclusively correlated with the presence of the 
setae but the majority of the those specimens 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



173 



that possessed setae on the arcessus also had long 
costal setae and the first Rs crossvein oblique. 
The possession of setae on the arcessus is unusual 
in Chrysopidae (Brooks & Barnard, 1990) and 
would be expected to be of taxonomic signifi- 
cance. However, Bram & Bickley (1963) found 
that of 12 males reared from a single female, one 
had setae on the arcessus and the other 11 
specimens had none which suggests this may be 
simply a case of intraspecific variation. 

Distribution. Canada, Mexico, USA. 

Material examined (41cf, 769) Bermuda, 
Canada, Mexico, USA: Arkansas, Florida, 
Georgia, Louisiana, Massachusetts, Mississippi, 
New York, North Carolina, Oklahoma, Oregon, 
Pennsylvania, South Carolina, Tennessee, 
Texas, Virginia. 



SYNOPTIC CHECKLIST OF THE 
SPECIES OF CHRYSOPERLA 

STEINMANN 

(* denotes types examined) 

adamsi Henry, Wells & Pupedis, 1993 [Nearctic] 

*annae sp. n. [Eastern Palaearctic] 

*asoralis (Banks, 1915) [Neotropics) 
*oscillans (Navas, 1922) syn. n. 
*lindana (Navsis, 1924) syn. n. 
*arequipae (Navas, 1929) syn. n. 

*carnea (Stephens, 1836) [Palaearctic] 
*affinis (Stephens, 1836) 
microcephala (Brauer, 1850) 
vulgaris (Schneider, 1851) 
lamproptera (Stein, 1863) 
*lucasina (Lacroix, 1912) syn. n. 
angelnina (Navas, 1931) 
ferganica (Navas, 1933) 
*pictavica (Lacroix, 1933) 
adaptata (Navas, 1934) syn. n. 
hmdbladi (Tjeder, 1939) 
maderensis (Tjeder, 1939) syn. n. 
canariensis (Tjeder, 1939) 
*shansiensis (Kuwayama, 1962) syn. n. 
nanceiensis Semeria, 1980 syn. n. 

*comanche (Banks, 1938) [Nearctic] 
sperryae (Banks, 1943) 

comans (Tjeder, 1966) [Afrotropics] 
*manselli Holzel, 1989 syn. n. 

*congrua (Walker, 1853) [Old World tropics] 
*concolor (Walker, 1853) 
otalatis (Banks, 1910) syn. n. 
*bequaerti (Navas, 1912) 



*lemoulii (Lacroix, 1923) syn. n. 
*clypealis (Navas, 1929) syn. n. 
solaria (Navas, 1930) 
*acutella (Navas, 1933) 

*decaryana (Navas, 1934) [Afrotropics] 

*defreilasi sp. n. [Neotropics] 

downesi (Smith, 1932) [Nearctic] 
mohave (Banks. 1938) syn. n. 

*dozieri (Smith, 1931) [Haiti] 

*exotera (Navas, 1914) [Neotropics] 
*extranea (Navas, 1922) syn. n. 
estradai (Navas, 1924) syn. n. 
*torrei (Navas. 1924) syn. n. 
'barbouri (Navas. 1924) syn. n. 
*tenera (NavSs. 1924) syn. n. 
exterior (Navas. 1925) 
cubensis (Navas, 1927) 
incisa (Banks, 1948) syn. n. 
quadornia (Banks, 1948) syn. n. 
castalia (Banks. 1948) syn. n. 

'externa (Hagen. 1861) [Neotropics] 
"lanata (Banks, 1910) 
graciana (Navas, 1919) 
*externa cocosensis Adams, 1983 syn. n. 

*exut (McLachlan, 1869) [St Helena] 
* ' wollastoni (Navas, 1914) 

*furcifera (Okamoto, 1914) [Eastern Palaearctic] 
savioi (Navas, 1933) syn. n. 
*kulingensis (Navas, 1936) syn. n. 

galapagoensis (Banks, 1924) [Galapagos Is] 

*gallagheri Holzel, 1989 [Oman] 

harrisii (Fitch, 1855) [Nearctic] 
stenosdgma (Navas, 1914) 

*insulata (Fraser, 1957) [Mascarene Is] 

*iranica (Holzel. 1967) [Iran] 

johnsoni Henry, Wells & Pupedis 1993 [Nearctic] 

mediterranea (Holzel, 1972) [Western Palaearctic] 

*mexicana sp. n. [Mexico] 

*mutata (McLachlan, 1898) [Western Palaearctic] 
*nepia (Navas, 1911) syn. n. 
*libera (Navas, 1928) syn. n. 
*quettana (Navas, 1931) syn. n. 
* phaeocephala (Navas, 1931) syn. n. 
*grazianii (Navas, 1932) syn. n. 
expurgata (Tjeder, 1949) 
*concinna (Holzel, 1973) syn. n. 

* nigrinervis sp. n. [Canary Is.] 

* nipponensis (Okamoto, 1914) stat. rev. (Eastern 
Palaearctic] 

kurisakiana (Okamoto, 1914) 

*ilota (Banks, 1915) syn. n. 



174 



S.J. BROOKS 



*kolthoffi (Navas, 1927) syn. n. 
*sinica (Tjeder, 1936) syn. n. 

*nyerina (Navas, 1933) [Afrotropics] 
*socia (Navas, 1936) syn. n. 

*oblita Holzel, 1973 [Eastern Palaearctic] 

*plicata (Tjeder, 1966) [Afrotropics] 

plorabunda (Fitch, 1855) [Nearctic] 
robertsonii (Fitch, 1855) 
pseudographa (Fitch, 1855) 
illinoiensis (Shimer, 1865) 
californica (Coquillet, 1890) syn. n. 

*pudica (Navas, 1914) [Afrotropics] 

incongrua (Navas, 1914) provisional syn. 
*nigriciana (Navas, 1931) syn. n. 

*renoni (Lacroix, 1933) [Western Palaearctic] 

* ankylopteryformis Monserrat & Diaz-Aranda, 
1989 syn. n. 

rufilabris (Burmeister, 1839) [Nearctic] 
interrupta (Schneider, 1851) 
*attenuata (Walker, 1853) 
*repleta (Walker, 1853) 
novaeboracensis (Fitch, 1855) 
tabida (Fitch, 1855) 
citri (Ashmead, 1880) 
medialis (Banks, 1903) 

*siamensis sp. n. [Thailand] 

*sillemi (Esben-Petersen, 1935) stat. rev. [Eastern 
Palaearctic] 

punensis (Ghosh, 1976) syn. n. 

gujaratensis (Ghosh, 1976) syn. n. 

sanandensis (Ghosh, 1977) syn. n. 

zastrowi (Esben-Petersen, 1928) [Afrotropics] 

NOT EXAMINED 

bellatula Yang & Yang, 1992 (BAU) 

chusanina (Navas, 1933) (ACAD SIN) 

hainanica Yang & Yang, 1992 (BAU) 

longicaudata Yang & Yang, 1992 (BAU) 

quinlingensis Yang & Yang, 1989 (BAU) 

sola Yang & Yang, 1992 (BAU) 

thelephora Yang & Yang, 1989 (BAU) 

xizangana (Yang & Yang, 1988) comb. n. (BAU) 

yulinica Yang & Yang, 1989 (BAU) 

SPECIES NEWLY EXCLUDED FROM CHRYSOP- 
ERLA 

Mallada anpingensis (Esben-Petersen) comb. n. 

Apertochrysa australis (New) comb. n. 

Mallada krakatauensis (Tsukaguchi) comb. n. 



Mallada maquilingi (Banks) comb. n. 
Mallada meloui (Navas) comb. n. 
Mallada satilota (Banks) comb. n. 
Mallada suzukii (Okamoto) comb. n. 
Apertochrysa triactinata (New) comb. n. 

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Figs 1-13 1-4, 6-13, Chrysoperla carnea; 5, C. sillemi. 1, fore wing; 2, fore wing f. typica (London, UK); 3, fore 
wing f. lucasina (Hertfordshire, UK); 4, hind tarsal claw (Spain); 5, hind tarsal claw (Pakistan); 6, hind tarsal 
claw (Crete); 7, hind tarsal claw (Israel); 8, apex d" abdomen lateral; 9, 0" gonarcus complex, dorsal; 10, C? 
gonarcus complex, lateral; 11, apex $ abdomen, lateral; 12, $ subgenitale, caudal; 13, $ spermatheca, lateral. 



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15 






Figs 14-23 14. 16, Chrysoperla downesi; 15, 21-22 C. plorabunda; 17, 19-20 C. carnea; 18, C. sillemi; 23, C. 
johnsoni. 14, fore wing; 15, hind tarsal claw; 16 hind tarsal claw; 17, O" tignum (Iran); 18, d* tignum (India); 19, 
O" tignum f. typica (London, UK); 20, C? tignum f. lucasina (Hertfordshire, UK); 21, cf tignum (Canada); 22, 
O" tignum (Tuscon, USA); 23 hind tarsal claw. 



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Figs 24-31 Chrysoperla zastrowi. 24, fore wing; 25, hind tarsal claw, lateral; 26, apex c? abdomen, lateral; 27, cf 
gonarcus complex, dorsal (gonosaccus not shown); 28, d" tignum, dorsal; 29, apex 9 abdomen, lateral; 30, 9 
subgenitale, caudal; 31, $ spermatheca, lateral. 



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Figs 32-39 Chrysoperla furcifera. 32, fore wing; 33, hind tarsal claw, lateral; 34, apex cf abdomen, lateral; 35, d* 
gonarcus complex, dorsal; 36, cf tignum, dorsal; 37, apex $ abdomen, lateral; 38, 9 subgenitale, caudal; 39,' $ 
spermatheca, lateral. 



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Figs 40-49 Chrysoperla iranica. 40, fore wing; 41. hind tarsal claw, lateral; 42, apex cf abdomen, lateral; 43, 
sternite 8+9, ventral; 44, a" tignum, dorsal; 45, cf gonarcus complex, lateral; 46, a" gonarcus complex, dorsal 
(gonosaccus not shown); 47, apex $ abdomen, lateral; 48, $ subgenitale, caudal; 49, $ spermatheca, lateral. 



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Figs 50-58 Chrysoperla mediterranea. 50, fore wing; 51, hind tarsal claw, lateral; 52, apex o" abdomen, lateral; 
53, cf tignum, dorsal; 54, cf gonarcus complex, dorsal; 55, O" gonarcus complex, lateral; 56, apex 9 abdomen, 
lateral; 57, $ subgenitale, caudal; 58. 9 spermatheca, lateral. 



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Figs 59-67 Chrysoperla nipponensis. 59, fore wing; 60, hind tarsal claw, lateral (Japan); 61, hind tarsal claw, 
lateral (Mongolia); 62, apex cf abdomen, lateral; 63, cf tignum, dorsal; 64, cf gonarcus complex, dorsal; 65, 
apex $ abdomen, lateral; 66, $ subgenitale, caudal; 67, 9 spermatheca, lateral. 



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Figs 68-75 Chrysoperla renoni. 68, fore wing; 69, hind tarsal claw, lateral; 70, apex O" abdomen, lateral; 71, c? 
gonarcus complex, dorsal; 72, cf tignum, dorsal; 73, apex 9 abdomen, lateral; 74, $ subgenitale, caudal; 75, $ 
spermatheca, lateral. 



186 



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Figs 76-83 Chrysoperla oblita. 76, fore wing; 77, hind tarsal claw, lateral; 78, apex d" abdomen, lateral; 79, ($ 
gonarcus complex, dorsal; 80, cf tignum, dorsal; 81, apex 9 abdomen, lateral; 82, 9 subgenitale, caudal; 83, 9 
spermatheca, lateral. 



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187 




Figs 84-89 Chrysoperla annae. 84, fore wing; 85, hind tarsal claw, lateral; 86, apex a" abdomen, lateral; 87, d" 
tignum, dorsal; 88, d* gonarcus complex, lateral; 89, d" gonarcus complex, dorsal. 



188 



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Figs 90-98 Chrysoperla gallagheri. 90, fore wing; 91, hind tarsal claw, lateral; 92, apex d" abdomen, lateral; 93, 
Cf gonarcus complex, dorsal (gonosaccus not shown); 94, cf gonarcus complex, lateral; 95, cf tignum, dorsal; 
96, apex $ abdomen, lateral; 97, 9 subgenitale, caudal; 98, 9 spermatheca, lateral. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



189 



100 x^II^ 




Figs 99-108 Chrysoperla comans. 99, fore wing; 100, hind tarsal claw, lateral; 101, apex (f abdomen, lateral; 
102, sternite 8+9 of cf abdomen, ventral; 103, cf gonarcus complex, lateral; 104, cT gonarcus complex, dorsal; 
105, cf tignum, dorsal; 106, apex 9 abdomen, lateral; 107, $ spermatheca, lateral; 108, 9 subgenitale, caudal. 



190 



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Figs 109-114 Chrysoperla decaryana. 109, fore wing; 110, hind tarsal claw, lateral; 111, apex c? abdomen, 
lateral; 112, cf tignum, dorsal; 113, ($ gonarcus complex, dorsal; 114, cf gonarcus complex, lateral. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



191 






Figs 115-123 Chrysoperla siamensis. 115, fore wing; 116, hind tarsal claw, lateral; 117, apex cS abdomen, lateral; 
118, cf gonarcus complex, lateral; 119, o" gonarcus complex, dorsal; 120, cf tignum, dorsal; 121, apex 9 
abdomen, lateral; 122, $ spermatheca, lateral; 123, 9 subgenitale, caudal (basal extension shown folded 
underneath subgenitale). 



192 



S.J. BROOKS 




Figs 124-132 Chrysoperla nyerina. 124, fore wing; 125, hind tarsal claw, lateral; 126, apex cf abdomen, lateral; 
127, cf tignum, dorsal; 128, cf gonarcus complex, dorsal; 129, cf gonarcus complex, lateral; 130, apex $ 
abdomen, lateral; 131, $ subgenitale, caudal; 132, 9 spermatheca, lateral. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



193 




Figs 133-141 Chrysoperla plicata. 133, fore wing; 134, hind tarsal claw, lateral; 135, apex d" abdomen, lateral; 
136, cf tignum, dorsal; 137, ($ gonarcus complex, dorsal; 138, a" gonarcus complex, lateral; 139, apex 9 
abdomen, lateral; 140, 9 subgenitale, caudal; 141, 9 spermatheca, lateral. 



194 



S.J. BROOKS 






Figs 142-148 Chrysoperla insulata. 142, fore wing; 143, hind tarsal claw, lateral; 144, apex c? abdomen, lateral; 
145, d" gonarcus complex, lateral; 146, apex 9 abdomen, lateral; 147, 9 subgenitale, caudal; 148, 9 
spermatheca, lateral. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



195 




Figs 149-159 Chrysoperla pudica. 149, fore wing; 150, hind tarsal claw, lateral; 152, apex cf abdomen, lateral; 
153, cf tignum, dorsal (Transvaal); 154, cf tignum, dorsal (Nigeria); 155, cf gonarcus complex, lateral; 156, cf 
gonarcus complex, dorsal; 157, apex 9 abdomen, lateral; 158, $ subgenitale, caudal; 159, $ spermatheca, 
lateral. 



196 



S.J. BROOKS 




Figs 160-170 Chrysoperla mutata. 160, fore wing; 161, hind tarsal claw, lateral (Punjab); 162, hind tarsal claw, 
lateral (Arabia); 163, hind tarsal claw, lateral (Algeria); 164, apex cf abdomen, lateral; 165, spinellae; 166, cf 
gonarcus complex, lateral; 167 cf tignum, dorsal; 168, apex £ abdomen, lateral; 169, $ subgenitale, caudal; 
170, 9 spermatheca, lateral. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



197 




Figs 171-178 Chrysoperla nigrinervis. 171, fore wing; 172, apex d" abdomen, lateral; 173, cf tignum, dorsal; 174, 
Cf gonarcus complex, lateral; 175, apex 9 abdomen, lateral; 176, $ subgenitale, caudal; 177, 9 spermatheca, 
lateral; 178 hind tarsal claw, lateral. 



198 



S.J. BROOKS 






Figs 179-187 Chrysoperla exul. 179, fore wing; 180, hind tarsal claw, lateral; 181, apex cf abdomen, lateral; 182, 
Cf gonarcus complex, lateral; 183, cf gonarcus complex, dorsal; 184, cf tignum, dorsal; 185, apex $ abdomen, 
lateral; 186, $ subgenitale, caudal; 187, $ spermatheca, lateral. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



199 




Figs 188-195 Chrysoperla congrua. 188, fore wing; 189, hind tarsal claw, lateral; 190, apex O" abdomen, lateral; 
191, cf tignum, dorsal; 192, cf gonarcus complex, dorsal; 193, apex 9 abdomen, lateral; 194, 9 subgenitale, 
caudal; 195, 9 spermatheca, lateral. 



200 



S.J. BROOKS 






Figs 196-202 Chrysoperla dozieri. 196, fore wing; 197, hind tarsal claw, lateral; 198, apex o" abdomen, lateral; 
199, cf tignum, dorsal; 200, d" apex of arcessus; 201, cf gonarcus complex, lateral (gonosaccus not shown); 202, 
gonarcus complex, dorsal (gonosaccus not shown). 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



201 



205 




Figs 203-208 Chrysoperla defreitasi. 203, fore wing; 204, apex cf abdomen, lateral; 205, d" gonarcus complex, 
dorsal; 206, cf gonarcus complex, lateral; 207, cf tignum, dorsal; 208, hind tarsal claw, lateral. 



202 



S.J. BROOKS 



209 




214 



Figs 209-217 Chrysoperla asoralis. 209, fore wing; 210, hind tarsal claw, lateral; 211, apex 9 abdomen, lateral; 
212, 9 spermatheca, lateral; 213, $ subgenitale, caudal; 214, apex cf abdomen, lateral; 215, cf tignum, dorsal; 
216, cf gonarcus complex, dorsal; 217, cf gonarcus complex, lateral. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



203 



219 




999 



Figs 218-226 Chrysoperla exotera. 218, fore wing; 219, hind tarsal claw, lateral; 220, apex o" abdomen, lateral; 
221, cT gonarcus complex, dorsal; 222, O" gonarcus complex, lateral; 223, d" tignum, dorsal; 224, apex $ 
abdomen, lateral; 225, 9 subgenitale, caudal; 226, 9 spermatheca, lateral. 



204 



S.J. BROOKS 



229 




Figs 227-236 Chrysoperla externa. 227, fore wing; 228, hind tarsal claw, lateral; 229, hind tarsal claw, lateral; 
230, apex d" abdomen, lateral; 231, c? tignum, dorsal; 232, o" gonarcus complex, dorsal; 233, d" apex of 
arcessus, dorsal; 234, apex 9 abdomen, lateral; 235, $ subgenitale, caudal; 236, 9 spermatheca, lateral. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



205 






Figs 237-245 Chrysoperla galapagoensis. 237, fore wing; 238, hind tarsal claw, lateral; 239, apex d" abdomen, 
lateral; 240, ($ gonarcus complex, caudal; 241, cf tignum, dorsal; 242, apex 9 abdomen, lateral; 243, 9 
subgenitale, caudal; 244, 9 subgenitale, ventral; 245, 9 spermatheca, lateral. 



206 



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Figs 246-255 Chrysoperla comanche. 246, fore wing; 247, hind tarsal claw, lateral; 248, apex cf abdomen, 
lateral; 249, cf tignum, dorsal; 250, cf gonarcus complex, dorsal; 251, cf gonarcus complex, lateral; 252, apex 
of arcessus; 253. apex $ abdomen, lateral; 254. 9 subgenitale, caudal; 255, 9 spermatheca, lateral. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



207 



256 





260 



Vy 265 




Figs 256-265 Chrysoperla harrisii. 256, fore wing; 257, hind tarsal claw, lateral; 258, apex cf abdomen, lateral; 
259, cf gonarcus complex, dorsal; 260, cf apex of arcessus; 261, cf gonarcus complex, lateral; 262, cf tignum, 
dorsal; 263, apex $ abdomen, lateral; 264, 9 spermatheca, lateral; 265, 9 subgenitale, caudal. 



208 



S.J. BROOKS 



266 




Figs 266-274 Chrysoperla mexicana. 266, fore wing; 267, hind tarsal claw, lateral; 268, apex cf abdomen, 
lateral; 269, cf tignum, dorsal; 270, cf gonarcus complex, dorsal; 271, cf apex of arcessus; 272, apex 9 
abdomen, lateral; 273, $ subgenitale, caudal; 274, 9 spermatheca, lateral. 



TAXONOMIC REVIEW OF COMMON GREEN LACEWING GENUS CHRYSOPERLA 



209 




Figs 275-285 Chrysoperla rufilabris. 275, fore wing; 276, hind tarsal claw, lateral (Texas, USA); 277, hind tarsal 
claw, lateral (Canada); 278, hind tarsal claw, lateral (North Carolina, USA); 279, apex (S abdomen, lateral; 
280, cf tignum, dorsal; 281, cf apex of arcessus; 282, o" gonarcus complex, dorsal (gonosaccus not shown); 283, 
apex 9 abdomen, lateral; 284, 9 spermatheca, lateral; 285, 9 subgenitale, caudal. 



210 



S.J. BROOKS 



INDEX 



This index is to species-group and genus-group names only. Principal references are in bold; synonyms are in italics. References to 

the key (p. 144) and the checklist (p. 173) are not included. 



acutella 165 
adamsi 142, 150 
adaptata 146. 147 
affinis 146 
Anachrysa 141 
angelnina 146 
ankylopteryformis 156 
annae 142, 157 
Anomalochrysa 141 
anpingensis (Mallada ) 144 
Apertochrysa 141, 142, 144 
arequipae 167 
asoralis 143. 167 
Atlantochrysa 141, 142 
attenuata 172 
australis (Apertochrysa) 144 

barbouri 167 
bellatula 144 
bequaerti 164 
boninensis (Mallada) 144 
brevicollis 161, 162 

calif or nica 150 

canariensis 146 

carnea 142, 146, 149, 152, 154, 156, 157 

castalia 168 

Chrysopidia 141 

Chrysotropia 141 

chusanina 155 

citri 172 

clypealis 164 

cocosensis 168, 169 

comanche 143, 168, 169, 170, 171 

comans 142, 158, 159 

concinna 163 

concolor 164 

congrua 143, 164, 165, 169 

cubensis 167 

Cunctochrysa 141 

decaryana 142, 158, 159 
defreitasi 143, 166 
desjardinsi (Mallada) 144 
Dichochrysa 141, 142, 144 
downesi 142, 151, 152, 172 
dozieri 143, 165, 166 

estradai 167 

exotera 143, 166, 167, 170 

expurgata 163 

exterior 167 

externa 143, 166, 168, 169, 170 

extranea 167 

exul 143, 164 

ferganica 146 



flava (Nineta) 147 
furcifera 142, 146, 152, 153 

galapagoensis 143. 169 
gallagheri 142, 157, 159 
graciana 168 
grazianii 163 
gujaratensis 149 

hainanica 144 
harrisii 143, 170, 171 

illinoiensis 150 

ilota 155 
incisa 167 
incongrua 161. 162 
inconspicua (Mallada) 144 
insulata 142. 161 
interrupla 171 
iranica 142, 153 
italotis (Plesiochrysa) 165 

johnsoni 142. 150, 172 

kolthoffi 148. 155. 156 
krakatauensis (Mallada) 144 
kulingensis 152 
kurisakiana 155 

lamproplera 146 
lanata 168 
lemouhi 164 
libera 162 
lindana 167 
longicaudata 144 
lucasina 146, 148 
lundbladi 146 

maderensis 146 

Mallada 141, 144 

manselli 158 

maquilingi (Mallada) 144 

medial is 172 

mediterranea 142, 146, 154, 156 

Meleoma 142 

meloui (Mallada) 144 

mexicana 143, 171 

microcephala 146 

mohave 150, 151. 152 

mutata 143. 158, 162, 163, 164 

nanceiensis 146 
nepia 162 
nigriciana 161 
nigrinervis 143, 163, 164 
nipponensis 142, 146. 155 
novaeboracensis 172 



nyerina 142. 158. 160. 161 

oblita 142. 146, 156, 157 
occipitalis 165 
oscillans 167 
otalatis 164 

Pcyerimhoffina 141 

phaeocephala 162 

pictavica 146 

plicata 142. 158. 160, 161 

plorabunda 142. 146, 150, 151 

pseudographa 150 

pudica 143. 158, 161, 162, 163 

punensis 149 

quadornia 167 
queitana 162 
quinlingensis 144 

renoni 142, 156 

repleta 172 

robertsonii 150 

rubella 165 

rufilabris 143. 167. 171. 172 

sanandensis 149 
satilota (Mallada) 144 
savioi 152, 153 
shansiensis 146, 147 
siamensis 142, 159 
sillemi 142. 149 
sinica 155 
socia 160 
sola 144 
solaria 165 
sperryae 170 
slenostigma 170 
Suarius 142 
suzukii (Mallada) 144 

tabida 172 

tenera 167 

thelephora 144 

torrei 167 

triactinata (Apertochrysa) 144 

vittata (Nineta) 147 
vulgaris 146 

wollastoni 164 

xizangana 144 

yulinica 144 

zastrowi 142, 146, 149, 152 



Bulletin of The Natural History Museum 
Entomology Series 

Earlier Entomology Bulletins are still in print. The following can be ordered from Intercept (address on inside 
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Volume 44 

No. 1 The taxonomy, biology and medical importance of Simulium amazonicum Goeldi (Diptera: 

Simuliidae), with a review of related species. A.J. Shelley, R.R. Pinger & M. A. P. Moraes. 1982. 

Pp. 1-30, 96 figs, 2 tables. £4.50 

No. 2 A revision of the genus Belonogaster de Saussure (Hymenoptera: Vespidae). O.W. Richards. 

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No. 3 The taxonomy and phylogeny of the genus Polyura Billberg (Lepidoptera: Nymphalidae). R.L. 

Smiles. 1982. Pp. 115-237, 159 figs, including 2 colour plates, 2 maps. £19.00 

No. 4 A taxonomic revision of the genus Gastrimargus Saussure (Orthoptera: Acrididae). J.M. Ritchie. 

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No. 1 A catalogue and reclassification of the Ichneumonidae (Hymenoptera) described by C.G. 

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No. 2 A taxonomic review of the genus Phlebolomus (Diptera: Psvchodidae). D.J. Lewis. 1982. 

Pp. 121-209, 38 figs, 13 maps. £12.60 

No. 3 Stenomine moths of the neotropical genus Timocratica (Oecophoridae). V.O. Becker. 1982. 

Pp. 211-306, 179 figs, 2 tables. £13.50 

No. 4 Afrotropical species of the myrmicine ant genera Cardiocondvla, Leptothorax , Melissotarsus, 

Messor and Cataulacus (Formicidae). B. Bolton. 1982. Pp. 307-370, 43 figs. £8.40 

Volume 46 

No. 1 The generic and tribal classification of spore-feeding Thysanoptera (Phlaeothripidae: 

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No. 2 A revision of the Afrotropical mole-crickets (Orthoptera: Gryllotalpidae). B.C. Townsend. 1982. 
Pp. 175-203, 63 figs. £4.20 

No. 3 Key to the genera of galerucine beetles of New Guinea, with a review of Sastra and related new 

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No. 4 The Afrotropical dacetine ants (Formicidae). B. Bolton. 1983. Pp. 267-416, 81 figs. £21.00 

Volume 47 

No. 1 A new genus of oriental lacewings (Neuroptera: Chrysopidae). S.J. Brooks. 1983. Pp. 1-26, 

95 figs. £4.00 

No. 2 The leafhopper genus Batracomorphus (Cicadellidae: Iassinae) in the eastern Oriental and 

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No. 3 The Afrotropical idiocerine leafhoppers (Homoptera: Cicadellidae). M.D. Webb. 1983. 

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Volume 48 

No. 1 Gelechiid moths of the genus Mirificarma. L.M. Pitkin. 1984. Pp. 1-70. 112 figs, 2 tables. £10.00 
No. 2 Macronematine caddisflies of the genus Atnphipsyche (Trichoptera: Hydropsychidae). P.C. 

Barnard. 1984. Pp. 71-130, 182 figs. £9.00 

No. 3 A review of the genera of Indo-Pacific Encyrtidae (Hymenoptera: Chalcidoidea). J.S. Noyes & 

M. Hayat. 1984. Pp. 131-395, 450 figs. £39.75 

Volume 49 

No. 1 Afrotropical jumping plant lice of the family Triozidae (Homoptera: Psylloidea-. D. Hollis. 1984. 

Pp. 1-102, 324 figs. £15.30 

No. 2 The taxonomy of the western European grasshoppers of the genus Euchorthippus, with special 

reference to their songs (Orthoptera: Acrididae). D.R. Ragge & W.J. Reynolds. 1984. 

Pp. 103-151, 88 figs. £7.20 

No. 3 An historical review of the higher classification of the Noctuidae (Lepidoptera). I.J. Kitching. 

1984. Pp. 153-234, 4 figs. £12.00 



No. 4 The Pimplinae, Xoridinae, Acaenitinae and Lycorininae (Hymenoptera: Ichneumonidae) of 

Australia. I.D. Gauld. 1984. Pp. 235-339, 100 figs, 18 maps. £15.75 

No. 5 The Palaearctic species of Ascogaster (Hymenoptera: Braconidae). T. Huddleston. 1984. 

Pp. 341-392, 79 figs. £7.80 

Volume 50 

No. 1 Taxonomy of Neotropical Derbidae in the new tribe Mysidiini (Homoptera). P.S. Broomfield. 

1985. Pp. 1-152, 501 figs. £22.80 

No. 2 Nymphal taxonomy and systematics of the Psylloidea (Homoptera). I.M. White & I.D. 

Hodkinson. 1985. Pp. 153-301, 201 figs, 18 tables. £23.00 

No. 3 The Whitefly of New Guinea (Homoptera: Aleyrodidae). J.H. Martin. 1985. Pp. 303-351, 

48 figs. £8.00 

Volume 51 

No. 1 The ichneumon-fly genus Banchus (Hymenoptera) in the Old World. M.G. Fitton. 1985. 

Pp. 1-60, 129 figs. £10.80 

No. 2 The phylogeny, classification and evolution of parasitic wasps of the subfamily Ophioninae 

(Ichneumonidae). I.D. Gauld. 1985. Pp. 61-185, 52 figs. £21.00 

No. 3 A cladistic analysis and classification of trichodectid mammal lice (Phthiraptera: Ischnocera). 

C.H.C. Lyal. 1985. Pp. 187-346, 250 figs. £26.00 

No. 4 The British and some other European Eriococcidae (Homoptera: Coccoidea). D.J. Williams. 

1985. Pp. 347-393, 18 figs. £8.00 

Volume 52 

No. 1 The sandflies of Egypt (Diptera: Phlebotominae). R.P. Lane. 1986. Pp. 1-35, 80 figs, 

4 tables. £5.60 

No. 2 Fungus moths: a review of the Scardiinae (Lepidoptera: Tineidae). G.S. Robinson. 1986. 

Pp. 37-181, 200 figs. £24.00 

No. 3 A revision of the European Agathidinae (Hymenoptera: Braconidae). G.E.J. Nixon. 1986. 

Pp. 183-242, 68 figs. £11.00 

No. 4 A key to the Afrotropical genera of Eucoilidae (Hymenoptera) with a revision of certain genera. 

J. Quinlan. 1986. Pp. 243-366, 359 figs. £21.00 

Volume 53 

No. 1 A review of Miletini (Lepidoptera: Lycaenidae). J.N. Eliot. 1986. Pp. 1-105, 108 figs. £18.00 
No. 2 Australian ichneumonids of the tribes Labenini and Poecilocryptini. I.D. Gauld & G.A. 

Holloway. 1986. Pp. 107-149, 65 figs. £8.40 

No. 3 The tribe Pseudophloeini (Hemiptera: Coreidae) in the Old World tropics with a discussion on 

the distribution of the Pseudophloeinae. W.R. Dolling. 1986. Pp. 151-212, 121 figs. £11.50 

No. 4 The songs of the western European grasshoppers of the genus Omocestus in relation to their 

taxonomy (Orthoptera: Acrididae). D.R. Ragge. 1986. Pp. 213-249, 128 figs. £7.50 

No. 5 The structure and affinities of the Hedyloidea: a new concept of the butterflies. M.J. Scoble. 

1986. Pp. 251-286, 102 figs. £7.00 

Volume 54 

No. 1 Studies on the Old World species of Holothrips (Thysanoptera: Phlaeothripidae). S. Okajima. 

1987. Pp. 1-74, 207 figs. £14.00 
No. 2 Spectacles and Silver Ys: a synthesis of the systematics, cladistics and biology of the Plusiinae 

(Lepidoptera: Noctuidae). I.J. Kitching. 1987. Pp. 75-261, 465 figs. £36.00 

No. 3 A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr 

(Hymenoptera: Formicidae). B. Bolton. 1987. Pp. 263-452, 100 figs. £36.50 

Volume 55 

No. 1 A reclassification of the European Tetrastichinae (Hymenoptera: Eulophidae), with a revision of 

certain genera. M.W.R. de V. Graham. 1987. Pp. 1-392, 744 figs. £75.00 

No. 2 The songs of the western European grasshoppers of the genus Stenobothrus in relation to their 

taxonomy (Orthoptera: Acrididae). D.R. Ragge. 1987. Pp. 393-424, 116 figs. £6.00 

Volume 56 

No. 1 The legume-feeding psyllids (Homoptera) of the wet Palaearctic Region. I.D. Hodkinson & D. 

Hollis. 1987. Pp. 1-86, 294 figs. £16.00 

No. 2 A review of the Malvales-feeding psyllid family Carsidaridae (Homoptera). D. Hollis. 1987. 

Pp. 87-127, 94 figs. £6.00 



No. 3 A review of the Rhadalinae ( = Aplocneminae) (Coleoptera: Melyridae). E.R. Peacock. 1987. 

Pp. 129-170, 59 figs. £8.00 

No. 4 A revision of some Afrotropical genera of Eucoilidae (Hymenoptera). J. Quinlan. 1988. 

Pp. 171-229, 199 figs. £11.00 

Volume 57 

No. 1 A survey of the Ophioninae (Hymenoptera: Ichneumonidae) of tropical Mesoamerica with 
special reference to the fauna of Costa Rica. I.D. Gauld. 1988. Pp. 1-309, 352 figs, 
32 maps. £52.00 

No. 2 A taxonomic revision of Alabagrus (Hymenoptera: Braconidae). M.J. Sharkey. 1988. 

Pp. 311-437, 28 figs, 22 maps. £24.50 

No. 3 A taxonomic revision of Caryocolum (Lepidoptera: Gelechiidae). P. Huemer. 1988. Pp. 439-571, 
221 figs. £25.00 

Volume 58 

No. 1 The mealybug genus Planococcus (Homoptera: Pseudococcidae). J.M. Cox. 1989. Pp. 1-78. 

40 figs. 
No. 2 The Simuliidae (Diptera) of the Santiago onchocerciasis focus of Ecuador. A.J. Shelley, M. 

Arzube & C.A. Couch. 1989. Pp. 79-130, 153 figs (including 2 plates in colour). 

Volume 59 

No. 1 The songs of the western European bush-crickets of the genus Platycleis in relation to their 
taxonomy (Orthoptera: Tettigoniidae). DR. Ragge. 1990. Pp. 1-35. 
A reclassification of the Melanotus group of genera (Coleoptera: Elateridae). C.M.F. von Hayek. 

1990. Pp. 37-115. 

No. 2 The green lacewings of the world: a generic review (Neuroptera: Chrysopidae). S.J. Brooks & 
P.C. Barnard. 1990. Pp. 117-286. 

Volume 60 

No. 1 The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini). PH. Williams. 

1991. Pp. 1-204. 

No. 2 Sattkria: a European genus of brachypterous alpine moths (Lepidoptera: Gelechiidae). L.M. 
Pitkin & K. Sattler. 1991. Pp. 205-241. 
A review of wing reduction in Lepidoptera. K. Sattler. 1991 . Pp. 243-288. 

Volume 61 

No. 1 Thrips (Thysanoptera) from Pakistan to the Pacific: a review. J.M. Palmer. 1992. Pp. 1-76. 
No. 2 Neotropical red-brown Ennominae in the genera Thysanopyga Herrich-Scaffer and Perissopteryx 
Warren (Lepidoptera: Geometridae). M. Kruger & M.J. Scoble. 1992. Pp. 77-148. 

Volume 62 

No. 1 Caloptilia leaf-miner moths (Gracillariidae) of South-East Asia. Decheng Yuan and Gaden S. 

Robinson. 1993. Pp. 1-37. 
No. 2 Neotropical Emerald moths of the genera Nemoria, Lissochlora and Chavarriella, with particular 

reference to the species of Costa Rica (Lepidoptera: Geometridae, Geometrinae). Linda M. 

Pitkin. 1993. Pp. 39-159. 

Volume 63 

No. 1 A revision of the Indo-Pacific species of Ooencyrtus (Hymenoptera: Encyrtidae), parasitoids of 
the immature stages of economically important insect species (mainly Hemiptera and 
Lepidoptera). D.W. Huang and J. S. Noyes. Pp. 1-135. 



'-■■ 



137 A taxonomic review of the common green lacewing genus Chrysoperla 
(Neuroptera: Chrysopidae) 

S.J. Brooks 



Natural his i oRY 

MUSEUM SHOPS 

139 -5^ 



ENTOMOLOGY SERIES 

Vol. 63, No. 2, November 1994